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Muelleria

Plant and Fungal Taxonomy & Systematics

REFERENCE

Vol 30(2) 2012

Royal _

Botanic

Gardens

Melbourne

Lastreopsis shepherdii (Kuntze ex Mett.)
Source: State Botanical Collection
Artist: Frederick Pitcher (1856-1935)

A founding member of the Field
Naturalists Club of Victoria, Pitcher trained
with Baron von Muelier specialising in
the Pteridophyta. He worked closely
with William Guilfoyle on the planting of
Australian species in Melbourne Botanic
Gardens. Using the nature printing
process, Pitcher made impressions of 60
species of ferns; some of which survive
in the Fern Gully today.

Royal Botanic Gardens
>.'-et bourse

9- JUL 2012
LIBRARY

Muelleria publishes research papers on Southern Hemisphere plant, alga! and fungal taxonomy and systematics,
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Muelleria

Contents

Volume 30(2) 2012

Royal Bo':aaic Gardens |
Melboiir.'ie j

3 - JUl 2Ci2

library

IU)ya] ^

Botanic

Gardens

Melbourne

Contributed papers Page

Five new endemic eucalypts for Victoria.,.83

- K. Rule

A revision of eastern Australian Bossiaea (Fabaceae: Bossiaeeae).106

- IR. Thompson

Genetic analysis suggests a wide regional provenance distribution for Epacris impressa .175

- M. Conomikes, GM. Moore, C McLean

Cyclosorus interruptus (Thelypteridaceae): new to Victoria.183

- S. Sinclair, V. Stajsic and G. Sutter

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Five new endemic eucalypts for Victoria

K. Rule

do National Herbarium of Victoria, Birdwood Avenue, South Yarra, Victoria, 3141, Australia.

Introduction

Throughout Victoria large tracts of eucalypt-dominated natural
vegetation have been lost to clearing for agriculture and other
commercial activities. In many parts of the state only remnant pockets
remain in protected reserves, on public land, including state forests and
roadside reserves, and on farms. In recent years extensive survey work
of accessible remnants of eucalypts has been undertaken by this author
and others, the result of which has been the discovery of several markedly
restricted, previously overlooked eucalypts worthy of taxonomic
consideration. Some new taxa have been described recently, some are
still under investigation and five are here given formal treatments as new
species.

All the new taxa are Victorian endemics and considered rare or
threatened.TheseareEt/co/ypfusbuny/p,atallish,slenderswampgumfrom
the Bunyip State Park in western Gippsland, £ conferto, a scentbark from
the Fryers Range near Castlemaine in north-central Victoria, £ caroloniae,
a mountain grey gum from Mt Martha on the Mornington Peninsula,
Eyarriambiack, a mallee-box from near Brim in the southern part of the
Victorian Mallee region, and £ aurifodina, a brown stringybark from the
Goldfields region of north-central Victoria (Fig. 1).

Taxonomy

Series Foveolatae {Swamp Gum Complex)

As the common name suggests, the swamp gums form a group of
eucalypts occurring in moist localities in south-east Queensland, New
South Wales, Victoria, Tasmania and south-east South Australia. In this
treatment 14 taxa, including the newly described £ bunyip, are included
in the key. £ ovata Labill. was described in 1806 and a large-fruited
form of the species, £ ovata var. grandifiora Maiden was described in
1916. Eucalyptus aquatica (Blakely) LA.SJohnson & K.D.Hill was first
described as a variety of £ ovata in 1934 but elevated to a species in
1990. Eucalyptus camphora R.T.Baker was described in 1899 and two
additional infraspecific taxa, namely subsp. humeana L.A.S.Johnson &
K.D.Hill and subsp. relicta LA.S.Johnson & K.D.Hill, were erected in 1990.

Abstract

Eucalyptus bunyip, a tall swamp gum
from the Bunyip State Park in west
Gippsland, £ conferta, a small-leaved
scentbark from the Fryers Range near
Castlemaine in north-central Victoria,

£ carolaniae, a rough-barked mountain
grey gum from Mt Martha on the
Mornington Peninsula, E.yarriambiack,
an umbrageous mallee-box from
near Brim in the southern part of the
Mallee region, and £ aurifodina, a
small-fruited brown stringybark from
the Victorian goldfields in the Avoca-
Castlemaine area, are described as
new Victorian endemic species. The
affinities, ecologies, distributions and
conservation statuses of each new
taxon are discussed.

Key words: short-range endemics,
swamp gum, scentbark, mallee-box,
mountain grey gum, stringybark

Muelleria 30(2): 83-105 (2012)

Ilotanic

CJardens

Muelleria

Rule

Figure 1 . Distribution map
for new species: Eucalyptus
bunyip (closed rectangle),
£ conferta (cross), £
carolaniae (closed circle),
£ yarriambiack (closed
triangle), £ aurifodina
fopen circle).

Eucalyptus aggregate H.Deane & Maiden was described
in 1900, £ rodwayii R.T.Baker & H.G.Sm. in 1912 and
£ yarraensis Maiden & Cambage in 1922. Eucalyptus
barberi LA.SJohnson & Blaxell was described in 1972,
£ brookeriana A.M.Gray in 1979, £ cadens J.D.Briggs
& Crisp in 1989 and £ strzeleckii Rule in 1992. Brooker
(2000) included £ macarthurii H.Deane & Maiden (1899)
in the Series Foveolatae. In my view, that species does
not belong with the swamp gums and may be better
placed elsewhere, possibly within the Series Vim'males
Blakely on the basis that its seedling morphology and
ontogeny are consistent with £ viminalis Labill. and its
subspecies.

The Series Foveolatae is characterised by the following:
The habit is usually a tree (small to tall) or rarely a mallee.
The bark in most species is smooth, often with a stocking
of accumulated loose strips, chunks or plates, or rough,
usually box-like, and extending to the major branches.
The Juvenile leaves are disjunct, petiolate and mostly
ovate. The adult leaves are ovate to lanceolate, green
and lustrous in most species. The inflorescences are
7-flowered or rarely 7-11-flowered.The buds are usually
pedicellate and diamond-shaped, or less often clavate
or ovoid. The fruits are pedicellate and mostly obconical.

The species described here as £ bunyip was first
brought to my attention bythelateJohn Reid and Neville
Walsh, both of MEL, whose survey work in the Bunyip
State Park led them to regard it as a distinctive entity.
Subsequent field studies and progeny trials endorsed
their initial assessment and, thus, it is described here as
a new species within the Series Foveolatae.

Eucalyptus bunyip Rule sp. nov.

Eucalypto strzeleckii affinis habitu elatiore graciliore,
foliis juvenilibus minoribus, foliis mediis persistentibus,
pedunculo delicato longiore, alabastris minoribus,
pedicellis longioribus, fructibus subcampanulatis
minoribus differt.

Type: Victoria: Camp Road, Bunyip State Park, on the
southern side of the creek crossing; 37° 59'07"S., 145°
38' 30" E., K. Rule 10507, 5.vi.2007. HOLD: MEL; ISO: AD,
CANB, NSW.

Trees slender, erect, to c. 40 m tall. Bark smooth, whitish
to light brown, becoming yellow-orange in spring; a
short, compact, corky dark grey stocking present at the
base. Seedling leaves ovate, at first sessile, becoming
shortly petiolate, slightly lustrous and green above, pale
green below. Juvenile leaves elliptical, ovate or ovate-
lanceolate, petiolate, apiculate, disjunct, discolorous,
green to blue-green, sub-lustrous above, whitish below,
4-6 cm long, 1.6-3 cm wide; margins entire; nodes
relatively remote; petioles 1-2 cm long. Intermediate
leaves broadly elliptical or broadly-ovate, rarely obovate,
dull or sub-lustrous, slightly discolorous or concolorous,
blue-green, to 8 cm long, 5.5 cm wide, persisting in large
numbers in the mature canopy. Coppice leaves with
lightly glaucous growth tips. Adult leaves ovate, ovate-
lanceolate or broadly lanceolate, slightly coriaceous,
acuminate, undulate, concolorous, densely reticulate,
sub-lustrous or lustrous, green, 10-17 cm long, 1.8-3.2
cm wide; nodes relatively remote; petioles 1.8-3.2
cm long; intramarginal vein 2-3 mm from margin; oil

Vol 30(2) 2012

Five new Victonan eucalypts

Key to the Swamp gums

1 Adult leaves dull, blue-green to glaucous. E, cadens

1: Adult leaves sub-lustrous or lustrous, green or slightly blue-green.2

2 Rough, box-like bark persistent to at least major branches. 3

2: Rough bark, if present, extending only to mid-trunk.5

3 Adult leaves ovate, to 3 cm wide.£ yarraensis

3: Adult leaves narrowly lanceolate or lanceolate, to 2 cm wide. 4

4 Intramarginal vein remote (to 3 mm from margin); peduncles to 4 mm long. E,aggregata

4: Intramarginal vein dose to margin (c 1 mm from margin); peduncles 5-8 mm long.£ rodwayi

5 Fruits 9-12 mm long, 9-12 mm diam.£ ovata var. grandiflora

5; Fruits 4-8 mm long, 4-8 mm diam.6

6 Oil glands sparse or apparently absent from adult leaves.£ ovata var. ovata

6: Adult leaves abundantly glandular. 7

7 Juvenile leaves crenulate; adult leaves discolorous.£ brookeriana

7: Juvenile leaves with entire margins; adult leaves concolorous.8

8 Mallees or small, slender trees.9

8: Robust, umbrageous trees or medium to tall upright trees.12

9 Adult leaves lanceolate; fruits more or less cupular. E. barberi

9: Adult leaves ovate or sub-orbicular; fruits obconical.10

10 Petioles of adult leaves > 2 cm long.£, camphora subsp. relicta

10; Petioles of adult leaves to 2 cm long. 11

11 Juvenile leaves to 5 cm long, 3 cm wide; petioles of juvenile leaves to 5 mm long;

petioles of adult leaves <1 cm long...£ aquatica

11: Juvenile leaves to 11 cm long, 5 cm wide; petioles of juvenile leaves > 2 cm long;

petioles of adult leaves 1-2 cm long. E, camphora subsp. camphora

12 Habit often an umbrageous tree, to 20 m tall; new season's growth tips non-pruinose;

juvenile leaves to 8 cm wide.£ camphora subsp. humeana

12: Erect trees to 40 m tall; new season's growth tips pruinose; juvenile leaves to 4 cm wide.13

13 Mature canopy a mixture of broadly ovate intermediate leaves and ovate or broadly

lanceolate adult leaves; pedicels equal to or longer than fruits; fruits 4-5 mm diam.£ bunyip

13: Mature canopy composed of ovate adult leaves; pedicels shorter than fruits; fruits 4-8 mm diam.£ strzeleckii

Muelleria

Rule

glands numerous, regular, island; new season's growth
tips glaucous. Inflorescences simple, axillary, 7-flowered;
peduncles slender and delicate, terete or angular,
9-14 mm long, c.1 mm thick, thickening slightly and
contracting when bearing mature fruits. Floral buds
clavate or slightly diamond-shaped, pedicellate, scarred,
often faintly pruinose in early development, 7-9 (-10)
mm long, 2.5-3.5 mm wide; pedicels slender, as long
as buds or to 1.2 times longer; hypanthium tapered;
operculum shortly rostrate, equal to or longer than the
hypanthium; locules 3 or 4; ovules in 4 vertical rows;
stamens irregularly flexed, all fertile; filaments white;
anthers dorsifixed, versatile, oblong, dehiscing through
longitudinal slits. Fruits sub-campanulate, distinctly
pedicellate, 5-6 mm long, 4-5 mm diam; pedicels
delicate and slender, longer than fruits, 6-10 mm long;
disc slightly elevated and rolled; valves usually slightly
exserted. Fertile seeds dark grey, elongated, flattened,
finely pitted; hilum ventral (Fig. 2).

Flowering Period: Autumn.

Distribution and Habitat: Eucalyptus bunyip occurs
in the narrow valleys of the Diamond and Black Snake
creeks in the Bunyip State Park approximately 60 km
to the east of Melbourne. Its habitat is highly specific,
being along the narrow valley floors which are subject to
seasonal inundation and prolonged impeded drainage.
The mean annual rainfall of the catchment which feeds
both the creeks exceeds 1200 mm (Fig. 1).

Additional specimens examined: VICTORIA: Bunyip State
Park, 30 m S of Gembrook-Tonimbuc Road, 0.9 km E of Camp
Road intersection, J.C. Reid 2195, 13j<i.1996 (MEL2109207);
100 m in a north-westerly direction from the intersection of the
Black Snake Creek Road andTowt Road, K. Rule 10707, 5.vi.2007
(MEL); Tonimbuc Road, 3.8 km from Tonimbuc Hall towards
Gembrook, K. Rule, 10307, 5.vi.2007 (MEL); Dyers Picnic Ground,
Black Snake Creek Road, K. Rule 10607, 5.vi.2007 (MEL).

Associated Species: E. bunyip grows in pure stands
and abuts forests of £ cypellocarpa L.A.SJohnson,

■nf

Figure 2. Eucalyptus bunyip (a) tree; (b) adult leaf; (c) seedling; (d) buds and fruits.

Vol 30(2) 2012

Five new Victorian eucalypts

£ obliqua L'Her. and a tallish form of £ ignorabilis
LA.SJohnson & K.D.Hill which is currently under study.
Other species occurring in the vicinity include £ dives
Schauer, £ cephalocarpa Blakely, £ radiata Sieber ex
DC. subsp. radiata, £ fulgens Rule, £ ovata var. ovata,
£ viminalis subsp. viminolis, and £ sieberi LA.SJohnson.

Conservation Status: Eucalyptus bunyip has a
distribution spread along several linear kilometres and
entirely within a protected reserve, the Bunyip State
Park, where an estimated 4000 mature trees exist.
According to lUCN criteria (lUCN 2001) a recommended
status for the species is 'vulnerable' (VU).

EtymologyiTUe epithet, of Aboriginal origin, is used
as a noun in apposition and refers to the Bunyip State
Park, the locality to which the species is restricted.

Discussion: Eucalyptus bunyip is distinguished within
the swamp gums by its tallish, slender habit, smooth
bark with a short stocking that is compact and corky,
its sub-lustrous green to blue-green, elliptical, ovate
or ovate-lanceolate juvenile leaves, its relatively broad,
blue-green intermediate leaves, which persist in the
inner canopy of mature trees, its abundantly glandular,
relatively narrow adult leaves, its lightly pruinose
immature buds and growth tips, its relatively long,
slender peduncles, its small, clavate to diamond-shaped
buds with a rostrate operculum and long, slender
pedicels and its small, sub-campanulate fruits. The
species is regarded as being a part of a narrow complex
within the swamp gums comprising £ camphora (and
its subspecies) and £ strzeleckii, whose features include

Table 1 . Comparisons between £ bunyip and related species.

Characters

£. strzeleckii

£. camphora subsp. humeana

£. bunyip

Habit

erect, robust trees to 40 m tall

usually umbrageous trees to

20 m tall

slender trees to 40 m tall

Bark

smooth

smooth with a box-like basal
stocking often extending to
mid-trunk

smooth with a short, compact,
corky basal stocking

Juvenile Leaves

Shape

lanceolate, ovate or ovate-
lanceolate

broadly ovate to orbicular,
often emarginate

elliptical, ovate or ovate-
lanceolate

Size

5-8 cm long, 1.6-4 cm wide

4-8 cm long, 4-8 cm wide

4-6 cm long, 1.6-3 cm wide

Colour, lustre

green to blue-green, sub-
lustrous

sub-glaucous, dull

green to blue-green, sub-
lustrous

Adult leaves

Shape

ovate

broadly ovate to sub-orbicular

broadly lanceolate to ovate

Size

8-20 cm long, 1.5-3 cm wide

8-15 cm long, 2.8-6.4 cm wide

10-17 cm long, 1.8-3.2 cm wide

Petiole length

1.6-3 cm long

2.2-4 cm long

1.8-3.2 cm long

Growth tips

pruinose

non-pruinose

pruinose

Canopy composition

adult leaves

inner canopy of Intermediate
leaves, outer canopy of adult
leaves

Peduncle length

7-14 mm long

10-18 mm long

9-14 mm long

Buds

Shape

ovoid or slightly diamond

diamond

clavate or slightly diamond

Size

5-8 mm long, 3-5 mm diam

5-7 mm long, 3-4 mm diam

7-10 mm long, c. 3 mm diam

Pedicel length

3-5 mm long

7-12 long

Operculum shape

shortly rostrate

conical

shortly rostrate

Fruits

Size

4-6 mm long, 5-8 mm diam

4-6 mm long, 4-6 mm diam

5-6 mm long, 4-5 diam

Shape

obconical to sub-campanulate

obconical

sub-campanulate

Pedicel length

2-4 mm long

6-10 mm long

Muelleria

Rule

adult leaves that possess an abundance of island glands.
Eucalyptus strzeleckii is considered to be its closest
relative, but it also shares features with E. camphora
subsp. humeana. Both of these taxa occur in the same
region as E bunyip (Table 1).

Eucalyptus strzelecki has a more extensive distribution
than E bunyip, occurring in south and west Gippsland
and in the western part of the Otway Ranges. Like
E bunyip, its growth tips are pruinose during the spring
growing period, it has similar juvenile leaves (except
those of E strzeleckii are longer), abundantly glandular
adult leaves and similar shaped fruits. However, it
differs from E bunyip by its preference for deep, fertile,
intermittently wet soils in a variety of sites, including
river banks, valley floors and slopes, rather than sites of
constantly impeded drainage preferred by E bunyip. It
also differs by its more robust habit with a stout trunk
and branches, the absence of the short stocking of
compact, corky bark, its mature canopy consisting
entirely of ovate-lanceolate or ovate adult leaves, its
shorter peduncles, its broader, more or less ovoid buds
with shorter pedicels and its generally larger fruits with
shorter pedicels.

Eucalyptus camphora subsp. humeana, also a relative
of E bunyip, is a widespread swamp gum occurring
along more or less elevated water courses of north-east
Victoria and adjacent areas of New South Wales. It is less
common in central and east Gippsland, but abundant in
the Yarra Valley which is about 20 km to the north-west
of the western-most population of E bunyip. Prior to this
study E bunyip had been regarded by local observers as
a forest form of E camphora The confusion between the
two was understandable as the mature canopies of both
contain relatively broad, richly glandular leaves and
have similar sized buds and fruits. However, the mature
canopy of E camphora subsp. humeana consists entirely
of broadly ovate or sub-orbicular adult leaves, whereas
E bunyip contains relatively broad intermediate leaves
in the inner canopy and much narrower ovate or broadly
lanceolate adult leaves at the ends of its branchlets.
Eucalyptus camphora subsp. humeana further differs
by its smaller, usually umbrageous habit (to 20 m tall),
its often substantial stocking of persistent, compact,
box-like bark, its larger, broadly ovate or orbicular, dull,
bluish Juvenile leaves, its broader, distinctly pendulous
adult leaves borne on longer petioles, Its non-pruinose
buds and new season's growth tips and its obconical
fruits borne on shorter, less delicate pedicels.

Series Acadiformes (Scentbark Complex)

The first descriptions for scentbarks, Series Acadiformes
Maiden, were for two species occurring in the New
England region of New South Wales. They were for
E ocaciiformis H.Deane & Maiden (1899) and E nicholii
Maiden & Blakely (1929). E aromaphloia L.D.Pryor
& J.H.Willis (1954) was described to accommodate
populations in central Victoria and E corticoso
LA.SJohnson (1962) for populations near Rylstone to
the north-west of Sydney and in the Grampians Ranges
of western Victoria. In 1971 Pryor and Johnson, placed
E corticosa, under E aromaphloia. It was not until
1989 that a study by Chappill etal. (1986) resurrected
E corticosa and identified three morphological and
geographical forms within the populations which had
been long-regarded as E aromaphloia (as well as the
typical form, forms for eastern Victoria and adjacent
areas of New South Wales and for the Victorian
Wimmera region). Eucalyptus ignorabilis L.A.SJohnson
& K.D.Hill was described in 1991 and E splendens
Rule (occurrences to the north-west of Portland),
E sobulosa Rule (populations in the Little Desert of the
Wimmera region and the Grampians) and E fulgens
Rule (populations in west Gippsland) were described In
1996. In the case of E sabulosa, Brooker and Slee (1996)
redefined it as a subspecies within E aromaphloia but
Nicolle (2006) regarded it a distinct species. E arcana
(D.Nicolle & Brooker) Rule (restricted to the Carpenter
Rocks area of Lower south-east South Australia) was first
described as a subspecies of E splendens in 2000 but
was elevated to a species in 2009. The classification of
both E splendens and E arcana have been somewhat
controversial. Both were placed within the Series
Viminales adjacent to E viminalis, the former by Brooker
and Slee (1997), and the latter by Nicolle and Brooker
(1998), but Nicolle (2006) regarded them as a part of
the Series Acadiformes. On the basis of their seedling
ontogeny being consistent with members of the Series
Acadiformes, despite being lustrous and green as in
E viminalis, they are regarded here as allies of that series
and thus are included in the key provided. In more
recent times further studies have shown that the small
but scattered population of scentbarks occurring in
the Fryers Range near Castlemaine in central Victoria is
sufficiently unique to be regarded as a separate species.
These populations are treated here as E conferta.

Vol 30(2) 2012

Five new Victorian eucalypts

Key for Scentbarks (Series Acadiformes)

1 Small, straggly trees with mature stems <20 cm diam; fruits tightly sessile. £, arcana

1: Robust trees with mature stems >50 cm diam; fruits pedicellate...„..2

2 Juvenile leaves lustrous and green; seedling stems square in cross-section and winged.E. splendens

2: Juvenile leaves dull, pale green, blue or glaucous; seedling stems neither square in cross-section nor winged.3

3 Juveniie leaves narrow (linear, narrow-elliptical, linear-lanceolate or narrow-lanceolate), <1 cm wide.4

3: Juvenile leaves broad (elliptical, elliptical-ovate or elliptical-lanceolate), 1.5-3 cm wide.7

4 Juvenile leaves pale green; seedling growth tips lustrous and green.E. sabulosa

4: Juvenile leaves blue-green or glaucous; seedling growth tips dull, blue-green or glaucous.5

5 Adult leaves to 1 cm wide; fruits with a flat disc. E.nicholii

5: Adult leaves to c 2 cm wide; fruits with an ascending disc.6

6 Juvenile leaves linear, falcate, crenulate; fruits 3-4 mm diam.E. conferta

6: Juvenile leaves linear-lanceolate or narrowly lanceolate; fruits 4-6 mm diam.E. corticosa

7 Outer rough bark dark brown; adult leaves lustrous and green; new season's growth tips bright green.E. fuigens

7: Outer rough bark light grey; adult leaves dull or sub-lustrous, blue-green or grey-green;

new season's growth tips blue-green to glaucous.8

8 Fruits hemispherical or slightly obconical; disc flat.E. acacUformis

8: Fruits ovoid or sub-globular; disc ascending.9

9 Adult leaves sub-lustrous, blue-green; fruits with short, thick pedicels.E. aromaphloia

9: Adult leaves dull, blue-grey; fruits with slender pedicels (often as long as fruits).E. ignorabilis

The adult features of taxa within the Series Acoc/Yfo/'mes
show only subtle differences in bark textures, adult
leaves (size, colour, lustre and oil gland density), and
fruits (shape and size and pedicel length and thickness),
which might suggest to some observers that some
entities, particularly those occurring in Victoria, would
be better placed as subspecies within f. aromaphloia.
However, to the contrary, progeny studies have shown
that the juvenile leaves of each contribute significantly
to its distinctiveness; glaucous, elliptical to ovate in
E. aromaphloia, sub-glaucous or glaucous, elliptical-
lanceolate to narrow-ovate and falcate in £ ignorablis,
pale green, lanceolate to narrow-ovate, often falcate
in £ fuigens, pale green, linear-elliptical and falcate in
£ sabulosa, lustrous, green, lanceolate or ovate-
lanceolate in £ splendens, lustrous, green, broadly ovate
in £ arcana, sub-glaucous, elliptical in £ acacUformis,

sub-glaucous, linear in £ nicholii, sub-glaucous, linear-
lanceolate in £ corticosa and glaucous, linear and falcate
in £ conferta.

Eucalyptus conferta Rule sp. nov.

Eucalypto aromaphloiae affinis habitu minore, foliis
juvenilibus angusto-falcatis subcrenatis confertis, foliis
adultis minoribus, pedunculo delicato, fructibus minoribus
differt.

Type: Victoria: Fryers Range, Vaughan Springs Road,
c 700 m S of intersection with Green Gully Road,
37°12'53" S., 144°14'32" E., K. Rule 02 W, 7.iv,2010. HOLO:
MEL; ISO: AD,CANB,NSW.

Smallish, slender frees, to c. 15 m tall. Bark grey-brown
or light brown, sub-fibrous, persistent to the upper
trunk, longitudinally furrowed, often loosely attached

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Rule

in short strips; bark on the lower trunk usually thick,
moderately furrowed, crusty; inner bark light brown;
upper bark decorticating in short ribbons; branches
smooth, whitish to pale brown. Seedling leaves elliptical,
sub-sessile, blue-green, discolorous; lower surface
white. Juvenile leaves linear, moderately to markedly
falcate, moderately crenulate, apiculate, sessile and
opposite, becoming shortly petiolate, disjunct, crowded
along the stem (not ericoid), slightly discolorous, dull,
glaucous, 4-8 cm long, 0.5-0.9 cm wide; petioles 0-5
mm long; growth tips lightly pruinose. Intermediate
leaves narrow-lanceolate or slightly ovate, often falcate.

crowded, petiolate, crenulate, slightly discolorous
to concolorous, dull, blue-green to glaucous. Adult
leaves narrow-lanceolate or lanceolate, sometimes
falcate, acuminate, moderately reticulate, dull, blue-
green or pale green, thin (0.18-0.27 mm thick),
7-11 (-13) cm long, 1.2-1.8 cm wide; petioles 1.2-1.8
cm long; intramarginal vein <2 mm from margin; oil
glands regular, numerous, mostly island; new season's
growth tips light green to sub-glaucous. Inflorescences
simple, axillary, 7-flowered; peduncles slender, slightly
angular, 7-11 mm long. Floral buds ovoid, pedicellate
scarred, 6-7 mm long, 2-3 mm wide; pedicels slender,

Figure 3. Eucalyptus conferta (a) foliage; (b) trunk with coppice leaves; (c) seedling; (d) buds; (e) fruits.

Vol 30(2) 2012

Five new Victorian eucalypts

2- 5 mm long;operculum conical, as wideashypanthium,

3- 4 mm long; stamens irregularly flexed, all fertile;
filaments white; anthers dorsifixed, oblong-cuneate,
versatile, dehiscing through longitudinal slits; locules
3 or 4; ovules in 4 vertical rows. Fruits slightly obconical
to sub-globoid, pedicellate, 3-4 mm long, 3-4 mm wide;
pedicels, slender, 1-3 mm long; disc ascending; valves
slightly exserted. Fertile seeds black, irregularly oblong,
flattened, lacunose; hilum ventral {Fig. 3).

Flowering Period: Autumn.

Distribution and Habitat: JUe new taxon is restricted
to the Glenluce State Forest in the Fryers Range, about
17 km to the south of Chewton (Fig. l).The population
is disjunct from other scentbarks, particularly
£ aromaphloiar which is known to occur approximately
20 km to the south in the Daylesford area. The species
occurs on hilly terrain and favours dry, shallow skeletal
soils that are sedimentary in origin. It is possible that
other stands of the taxon occur on similar sites in the
district. The mean annual rainfall of nearby Castlemaine
is approximately 550 mm, most of which falls in winter.

Additional specimens examined: VICTORIA: c. 5 km SE
of Vaughan, 0.5 km S of Sebastopol Ck., 10 m E of road on
Crown land, B. Kemps.n., n.v.l986 (MEL686536); Fryers Range,
Vaughan Springs Road c. 100 m N of intersection with Green
Gully Road, K, Rule 11007, 25.vi.2007 (MEL); Fryers Range,
Vaughan Springs Road, at the end of the bitumen, K. Rule 11207,
25.vi.2007 (MEL); Fryers Range, Green Gully Road, 700 m from
Vaughan Springs Road, K. Rule 11407, 25.vi.2007 (MEL); Fryers
Range, Green Gully Road, 1.1 km from Vaughan Springs Road,
K. Rule 11507. 25.vi.2007 (MEL).

Associated Species: Eucalyptus dives, E. nortonii
(Blakely) L.A.SJohnson,£.me///odoraA.Cunn.ex Schauer,
£ polyanthemos Schauer subsp. vestita L.A.SJohnson &
K.D.Hill, £ macrorhyncha F.Muell. ex Benth. and £ obliqua
all occur within the range of £ conferta.

Etymology: The name is from the Latin confertus
'crowded' in reference to the numerous pairs of leaves
occurring along the axis in the juvenile and intermediate
stages of development.

Conservation Status: The number of mature trees of
£ conferta is estimated to be between 400 and 500. Even
though theyoccurin a state forest, which hasnoapparent
value for timber extraction, its long-term security is not
guaranteed. In accordance with lUCN criteria (lUCN
2001) a status ofVulnerable'(VU) is suggested.

Discussion: Eucalyptus conferta is distinctive by its
smallish, sometimes spindly habit, its often loosely

attached, usually crusty lower bark, its linear, falcate,
glaucous, crenulate juvenile leaves, which become
crowded in the advanced seedling stage, its smallish,
thin, dull, blue-green to sub-glaucous adult leaves, its
delicate peduncles and its small buds and small, sub-
globular fruits that are borne on short, delicate pedicels.

Eucalyptus conferta is regarded as being closely related
to £ aromaphloia which differs by its generally larger
habit (stout-trunked, spreading trees), its thicker, more
deeply furrowed bark, its broader, ovate to elliptical,
non-falcate, non-crenulate, less crowded juvenile leaves,
its more glossy, thicker, larger adult leaves, its generally
shorter peduncles, its generally thicker, shorter pedicels
and its larger buds and fruits (Table 2).

The narrow juvenile leaves of £ conferta resemble
those of £ nicholii, a species from northern New South
Wales, as do its delicate peduncles and pedicels.
However, the appearance of its canopy differs from that
species by its generally broader adult leaves. Further,
the fruits of £ conferta are generally larger and more
globoid than those of £ nicholH and have a shape similar
to other southern scentbarks including £ aromaphloia,
£ sabulosa, £ ignorabilis and £ fulgens. Similarly, it shares
features with £ corticosa, namely its general appearance,
particularly its mature canopy and its delicate peduncles
and pedicels. However, £ corticosa differs from the new
species by its broader, non-falcate, non-crenulate, less
crowded juvenile leaves and its larger buds and fruits.

Series Globulares (Mountain Grey Gum
Complex)

In recent years several taxa regarded as mountain
grey gums have been described. The complex features
medium to tall or less often small trees, smooth bark
or box-type bark on the lower trunk, seedling stems
usually square in cross-section and sometimes winged,
juvenile leaves usually opposite, sessile and amplexicaul
for numerous pairs, green and glossy or glaucous in one
species, long, glossy and green adult leaves, long, strap¬
like peduncles, pedicellate, usually cigar-shaped buds
that are waisted at the mid-point and elongated fruits
with the valves enclosed. This group is closely related to
the long-leaf boxes (£ goniocalyx F. MuelI.ex Miq. and its
allies) which usually feature species with smaller habits,
box-type bark over most of the trunk, orbicular, usually
glaucous juvenile leaves, generally shorter adult leaves,
shorter peduncles and mostly sessile buds and fruits.

Muelleria

Rule

Table 2, Comparisons between £ conferta and £ oromaphloio

Characters

E, aromaphloia

E. conferta

Habit

umbrageous tree to 25 m tall

slender tree to 15 m tall

Bark

sub-fibrous, thick, deeply furrowed

sub-fibrous, moderately furrowed

Juvenile leaves

Size

2-6 cm long, 1 -4 cm wide

4-8 cm long, 0.5-0.9 cm wide

Shape

elliptical to ovate

linear, falcate

Colour

glaucous

Margins

entire

moderately crenulate

Density along the axis

not crowded

crowded

Adult leaves

Shape

lanceolate

Size

8-18 cm long, 1 -2.5 cm wide

7-13 cm long, 1.2-1.8 cm wide

Petiole length

1.2-2.2 mm

1.2-1.8 mm

Colour, lustre

blue-green, sub-lustrous

blue-green or pale green, dull

Thickness

0.24-0.39 mm

0.18-0.27 mm

Peduncle

Length

4-9 mm

7-11 mm

Buds

Size

4-7 mm long, 3 mm diam

3-5 mm long, 2-3 mm diam

Pedicel length

1-3 mm long

2-5 mm long

Fruits

Shape

hemispherical to sub-globular

sub-globular

Size

4-7 mm long, 4-7 mm diam

3-4 mm long, 3-4 mm diam

Pedicel length

0-2 mm long

1-3 mm long

The name £ goniocalyx was previously incorrectly
applied to tall forest trees which were named
£ cypellocarpa in 1962. Until recently all known
populations of mountain grey gums occurring in coastal
and sub-coastal regions from northern New South Wales
to western Victoria, were included with £ cypellocarpa.
Eucalyptus alaticaulis RJ.Watson & Ladiges (1987) was
described to cater for populations occurring in the
Grampians RangesofwesternVictoriaand near Anglesea
to the south-west of Melbourne. Eucalyptus retinens
LA.SJohnson & K.D.Hill, £ volcanica LASJohnson &
K.D.Hill (both 1990), £ oresb/a J.T.Hunter & JJ.Bruhl and
£ quinniorum J.T.Hunter & JJ.Bruhl (both 1999) are
names applied to mountain grey gums occurring in
northern New South Wales. Eucalyptus litoralis Rule
was described to accommodate the Anglesea 'form' of
£ alaticaulis and £ pyrenea Rule to cater for the disjunct
occurrences of mountain grey gums on Mt Avoca in the
Pyrenees Range of west-central Victoria (Rule 2004).

The mountain grey gum complex has been marked
by controversy as there is no consensus regarding the
acceptance of several of the taxa. In fact, Brooker and
Slee (1997) did not accept £ alaticaulis and Brooker
and Kleinig (2006) rejected £ retinens, £ oresbia,
£ quinniorum and £ pyrenea as distinct taxa, all of which
were regarded as synonymous with £ cypellocarpa.
Nicolle (2006) chose to accept £ alaticaulis as a distinct
taxon, but regarded £ pyrenea as a form of that species.
He further regarded £ oresbia and £ quinniorum as
forms of £ cypellocarpa. On the other hand, Hill (2002)
accepted all taxa occurring in New South Wales whilst
all Victorian taxa, including £ alaticaulis and £ pyrenea,
were accepted by Walsh and Stajsic (2007). Obviously, a
formal revision of the complex is required to eliminate
this controversy. At this point, however, all taxa listed
above are accepted and included in the key.

An additional taxon, which has long been regarded as
a form of £ goniocalyx, is not only treated here as a new

Vol 30(2) 2012

Five new Victorian eucalypts

Key to the Mountain Grey Gums

1 Juvenile leaves lanceolate (narrow to broad). 2

1: Juvenile leaves elliptical or ovate to orbicular.3

2 Mallees or rarely small, single-stemmed trees to 16 m tall; buds with operculum wider than hypanthium. E. quinniorum

2: Single-stemmed trees to 60 m tall; buds with operculum narrower than or as wide as the hypanthium.f. cypetlocarpa

3 Bark smooth throughout; buds not waisted at the mid-point.£, oresbia

3: Rough bark persistent on lower trunk or higher; buds waisted at the mid-point.4

4 Juvenile leaves glaucous. e, volcanica

4: Juvenile leaves green or blue-green. 5

5 Seedling stems without prominent wings. 6

5: Seedling stems prominently winged. 7

6 Longest adult leaves to 29 cm long. f. carolaniae

6; Longest adult leaves to 18 cm long. E.retinens

1 Fruits 5-6 mm diam.E pyrenea

7: Fruits 7-11 mm diam.8

8 Operculum 4-8 mm long; fruits 9-13 mm long, 8-11 mm diam.E litoralis

8: Operculum 3-4 mm long; fruits 6-10 mm long, 6-9 mm diam.E alaticaulis

species, but is regarded as belonging to the mountain
grey gum complex on the basis of both seedling and
adult characters. It occurs on the Mornington Peninsula
to the south-east of Melbourne and was first brought
to my attention in the early 1990s by the late Ms. Pat
Carolan, after whom it is named.

Eucalyptus carolaniae Rule sp. nov.

Eucalypto cypellocarpae affinis habitu minore, cortice
persistenti buxiformi, foliis juvenHibus latioribus griseo-
viridibus, foliis adultis majoribus, fructibus sessilibus vel
subsessilibus differt

Type; Victoria: Norfolk-Hopetoun Reserve, Mt Martha,
38°16'S. ]45°0VE.,PCarolan s.n., 9.vi.1989. HOLO: MEL
(3 sheets: MEL 117392, MEL 117393, MEL 117394).

Small-medium, robust, often spreading trees to
20 m tall. Bark sub-fibrous, box-like, grey-brown, deeply
furrowed, often crusty, persisting on lower trunk or
higher, graduating to thin, finely furrowed bark above;
branches usually smooth, pale grey-brown; old bark

decorticating in ribbons. Seedling leaves broadly ovate,
discolorous, pale green, sub-lustrous on the upper
surface, whitish on the lower surface. Seedling stems
initially square in cross section, becoming round by
the 6*'’ node. Juvenile leaves broadly ovate to broadly
elliptical or sub-orbicular, opposite, sessile, amplexicaul
for numerous pairs, apiculate, discolorous, lustrous and
light green, becoming dull or sub-lustrous and blue-
green with age, 6-10 cm long, 4-7 cm wide; growth
tips faintly glaucous throughout the seedling stage.
Intermediate leaves broadly ovate or ovate-lanceolate,
shortly petiolate, disjunct, lustrous, green to blue-green,
longer than Juvenile leaves. Adult leaves lanceolate,
falcate, pendulous, acuminate, concolorous, lustrous,
green, 16-29 cm long, 2-4 cm wide; petioles 1.7-3.2
cm long; venation conspicuous, moderately dense;
intramarginal vein remote, 3-5 mm from the margin;
glands regular, scattered, island and intersectional.
Inflorescences simple, axillary, 7-flowered; peduncles
flattened, thickening with age, 12-20 mm long,
3-4 mm wide. Floral buds ovoid-cylindrical, waisted at

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Rule

mid-point, scarred (outer operculum shed in early bud
development), ribbed, shortly pedicellate, 7-10 mm
long, 3-5 mm wide; pedicels 1-3 mm long; operculum
sharply conical 3-4 mm long, flush with the hypanthium
at the abscission zone; stamens inflexed; filaments
white, all fertile; anthers oblong, sub-basifixed, versatile,
dehiscing through longitudinal slits; locules (3) 4,
each with 4 ovular rows. Fruits cupular, sub-cylindrical
or slightly obconical, basally-tapered, sessile or sub-
sessile, moderately or prominently ribbed, thin-walled.

8-11 mm long, 6-9 mm diam, pedicels 0-2 mm long;
disc descending; valves (3-) 4, enclosed. Fertile seeds
dark brown or black, flattened, ovoid, 2-3 mm long,
tapered at one end, lacunose; hilum ventral (Fig. 4).

Flowering Period: Summer,

Distribution and Habitat The new taxon is known
only from a single population located above Mt Martha
township on the Mornington Peninsula approximately
60 km south-south-east of Melbourne (Fig. 1). It grows
along a sheltered gully in deep soils derived from granite.

Figure 4. Eucalyptus carolaniae (a, b) trees; (c) bark; (d) juvenile leaves; (e) fruits.

Vol 30(2) 2012

Five new Victorian eucalypts

The mean annual rainfall of the area is approximately
650 mm, most of which falls in winter.

Additional specimens examined: VICTORIA: Norfolk-
Hopetoun Reserve, Mt Martha, K. Rule 0201, 20.i.2001 (MEL);
Mt Martha Golf Course, K. Rule 0252, 15.V.2002 (AD, CANB, MEL,
NSW).

Associated Species: Eucalyptus radiata subsp. radiota,
E. viminalis subsp. viminalis and E. ovata var. ovata are
sympatric with £ carolaniae, whilst £ obliqua, and
£ pauciflora Sieber ex Spreng. subsp. pauciflora occur
in the vicinity. Hybrids with £ viminalis have been
observed in the field.

Etymology: The epithet honours the late Ms Pat
Carolan who collected the type specimen of the new
taxon and who first brought the existence of the
population to my attention. She is further recognised
for her contributions to the understanding of Victoria's
eucalypts, having been an enthusiastic collector and
having lodged numerous specimens with MEL

Conservation Status: An ecological study by Picone
and McCaffrey (2006) estimated the population of
£ carolaniae as approximately 500 naturally occurring
trees and saplings, nearly all of which occur along a
linear strip of bushland forming the Norfolk-Hopetoun
Reserve and extending on to a section of the lower
extremity of the Mt Martha Golf Course. A large
proportion of the population is sandwiched along a
narrow easement between housing blocks between
Norfolk and Hopetoun Roads. The habitat until recently
has been infested with blackberries and other weeds.
Its close proximity to the golf course and houses
remains a concern as the long term impact of changes
to nutrient levels and to drainage patterns is unknown.
A revegetation program has reintroduced several
hundred seedlings to an area of the golf course which
abuts naturally occurring trees,The reserve and the golf
course, which are managed by the Mornington Shire
Council, offer some protection to the species. As the
taxon's numbers are critically low and there are threats
from changes to drainage patterns and nutrient levels,
in accordance with lUCN criteria (lUCN 2001) a status of
'endangered'(EN) is considered appropriate.

Discussion: The combination of features which
distinguish £ carolaniae are its relatively small habit,
its rough, box-type bark which usually extends to the
upper trunk and sometimes to the major branches, its
seedling stems that are square in cross-section in early

seedling development, its Juvenile leaves that are ovate
to broadly elliptical or sub-orbicular and initially lustrous
and light green but become dull and blue-green with
age, its large adult leaves, its long, flattened peduncles,
its shortly pedicellate, cigar-shaped buds and its basally-
tapered, sessile or sub-sessile, thin-walled fruits.

Eucalyptus carolaniae shares features with both
£ cypellocarpa and £ goniocalyx which may raise the
possibility that it is derived from hybrid stock. Initially
the proposition of hybridism had some merit as the
adult trees exhibit slight variation in the amount of
persistent rough bark on the trunk. Despite this, other
adult features within the population are uniform and
repeated seedling trials have consistently produced
uniform seedlings, all of which supports £ carolaniae as
as a true-breeding, distinct species (Table 3).

Eucalyptus cypellocarpa differs from £ carolaniae by its
taller, straighter habit, its smooth bark, except for strips
and plates of old bark often remaining attached to the
base or iower trunk, its more lustrous, greener, narrower
(lanceolate) juvenile leaves, its generally shorter,
narrower adult leaves and its consistently pedicellate
buds and fruits.

Eucalyptus carolaniae resembles £ goniocalyx in habit
and bark and the type specimens were initially held
under that name at MEL. However, it is here regarded
as being a part of the mountain grey gum complex
and closely related to £ cypellocarpa. It differs from
£ goniocalyx by its seedling stems being initially square
in cross-section in early seedling development (stems
round in cross-section in £ goniocalyx), its broadly ovate
to broadly elliptical, apiculate juvenile leaves (seedlings
with some orbicular and emarginate juvenile leaves in
£gon/oca/yx),itslongeradultIeaves, itslongerpeduncles.
Its shortly pedicellate buds and its consistently basally-
tapered fruits.

Series Buxeales (The Mallee-boxes)

A new box species, £ yarriombiack, is described below.
It has close affinities with several other box species,
some of which, until recently, had been referred to as
mallee-boxes and placed in the Series Subbuxeales
Blakely (1934) by various commentators, for example,
Chippendale (1988), Brooker and Slee (1997), Nicolle
(2000) and Rule (2004). Such species include £ odorata
Behr, £ polybractea R.T.Baker, £ viridis R.T.Baker,

Muelleria

Rule

Table 3. Comparisons between E carolaniae and related species. Included in the key are references to various subspecies of
E. goniocalyx, three of which were described by Rule (2011).

Characters

£ goniocalyx

£. cypellocarpa

E. carolaniae

Habit

single or few-stemmed trees to 15 m
tall or mallees in subsp. exposa, to

25 m in subsp. fo//ax

trees to 60 m tall

single or few-stemmed trees
to 20 m tall

Bark

box-type to minor branches or smooth
in subsp. exposa, non-fractured in
subsp. fallax, loosely attached in
subsp. laxa

smooth except for old strips
and plates on lower trunk

usually box-type to upper
trunk

Seedling stems

round In cross-section

square in cross-section in
Victorian populations

square in cross-section for a
few pairs

Juvenile leaves

Shape

broadly ovate to orbicular

lanceolate

broadly ovate to broadly
elliptical or sub-orbicular

Apex

apiculate and emarginate

apiculate

Size

14-11 cm long, 4-10 cm wide

10-17 cm long, 3-7 cm wide

6-10 cm long, 4-7 cm wide

Colour, lustre

blue-green to glaucous, dull or green,
lustrous in subsp. viridissima

green, lustrous

green or blue-green, sub-
lustrous

Adult leaves

Size

8-20cm long, 1.5-3.5 wide, to 25 cm
long in subsp./flxa

11 -20 cm long, 1 -2.5 cm wide

16-29 cm long, 2-4 cm wide

Petiole length

1.6-3.3 cm

1.5-2.7 cm

1.7-3.2 cm

Peduncle length

6-15 mm long, to 18 mm long in
subsp. fallax

8-22 mm long

12-20 mm long

Buds

Shape

ovoid to cylindrical

Size

8-13 mm long, 3-6 mm diam

7-11 mm long, 3-5 mm diam

7-10 mm long, 3-5 mm diam

Pedicel length

sessile, rarely sub-sessile in subsp. laxa

2-3 mm

1-3 mm

Fruits

Shape

cupular or cylindrical

cupular to barrel-shaped

cupular or slightly obconical

Size

5-10 mm long, 5-10 mm diam

5-10 mm long, 6-9 mm diam

8-11 mm long, 6-9 mm diam

Pedicel length

sessile, rarely sub- sessile in subsp. laxa

1-2 mm

0-2 mm

£ wimmerensis Rule, £ aenea K.D.Hill, £ castrensis K.D.Hill,
£ walshii Rule, £ filiformis Rule and £ hawkeri Rule.Their
shared features include a mallee or small, tree-like habit,
either smooth bark throughout or persistent box-like
bark present to various heights on the stems, adult
leaves with numerous large, irregular island glands,
inflorescences that are simple and axillary, buds with
the outer operculum intact at anthesis and fruits that
are 3- or4-loculed.

Brooker (2000) in his revision of Eucalyptus discarded
the Series Subbuxeales and assigned the mallee-box
species to the Supraspecies Odoratae within the Series
Buxeales. In the key provided, this treatment only
partly adheres to Brooker's classification as it excludes

some species contained in his Supraspecies Odoratae
(£ albopurpurea (Boomsma) D.Nicolle and £ persistens
LASJohnson & K.D.Hill subsp. persistens, including
£ persistens subsp. tardecidens L.A.S.Johnson & K.D.Hill)
and other species that have been previously regarded
as mallee-boxes (£ porosa F.Muell. ex Miq., £ bosistoiana
F.Muell., £ froggattii Blakely and £ silvestris Rule) as they
do not comply totally with the criteria for the mallee-
box complex set out above. Eucalyptus albopurpurea has
adult leaves with a dense reticulation with intersectional
glands and paniculate inflorescences. Both subspecies
of £ persistens have paniculate inflorescences.
£ porosa has 5- or 6-loculed fruits and a uniquely remote
intramarginal vein. £uca/yprus bos/sfo/ano has paniculate

Vol 30(2) 2012

Five new Victorian eucalypts

Key to £. odorata and its allied Mallee-boxes

1 Juvenile leaves <0.6 cm wide.....2

1: Juvenile leaves 0.7-3,2 cm wide..... 3

2 Adult leaves lustrous and green; coppice leaves green.f. viridis

2: Adult leaves sub-lustrous and blue-green; coppice leaves glaucous.£ filiformis

3 Bark smooth throughout or rough bark as a short basal stocking on larger stems.4

3: Rough bark extending to upper stem or higher.8

4 Seedlings and buds pruinose.£. polybractea

4; Seedlings and buds non-pruinose.. 5

5 Juvenile leaves lustrous, green; fruits 2-4 mm diam...E aenea

5: Juvenile leaves dull, blue-green; fruits 4-6 mm diam.6

6 Slender, pole-like trees.E. walshii

6:Mallees...7

7 Juvenile leaves 1-1.5 cm wide; adult leaves dull or sub-lustrous, blue-green. E. wimmerensis

7: Juvenile leaves 1.5-4 cm wide; adult leaves lustrous, green.£. castrensis

8 Mallees; juvenile leaves glaucous; immature fruits pruinose.E polybractea

8:Trees; juvenile leaves green or blue-green; immature fruits non-pruinose.9

9 Slender, single or few-stemmed erect trees; adult leaves weeping; new season's growth tips lustrous, green.E bawkeri

9: Robust, few-stemmed, spreading trees: adult leaves erect; new season's growth tips dull, blue-green or sub-glaucous.10

10 Buds lightly pruinose; fruits 3-4 mm diam. E, yarriambiack

10: Buds non-pruinose; fruits 4-7 mm diam.E. odorata

inflorescences and 5- or 6-loculed fruits. Eucalyptus
froggattii has paniculate inflorescences and lateral veins
that do not connect with the intramarginal vein. Lastly,
£ silvestris has adult leaves with a dense reticulation and
sparsely distributed intersectional glands and in this
respect is allied to £ microcarpa Maiden.

Eucalyptus yarriambiack Rule sp. nov.

Eucalypto wimmerensi offinis habitu arborescent!
majore, cortice buxlforme, foliis juvenlHbus viridibus,
alobastris subpruinosis, fructibus minoribus differt et
Eucalypto odorata affinls foliis juvenillbus ongustioribus
viridibus minoribus, foliis adultis angustioribus
subnitentibus, alobastris subpruinosis minoribus, fructibus
minoribus differt.

Type: Victoria: Henty Highway, 1.6 km N of Brim, 36°
03'41"S., 142° 25' 13"E., K. Rule 2605, 18.iii.2005. HOLO:
MEL

Robust, spreading small trees, usually with a few thick
trunks, to 10 m tall, 10 m wide. Bark light grey-brown, box
type, persisting to at least to the upper trunk, smooth,
light grey-brown above; old bark decorticating in broad
strips. Seedling leaves narrowly elliptical, petiolate,
opposite for 3 or 4 pairs, green, discolorous. Juvenile
leaves narrowly lanceolate or elliptical-lanceolate,
alternate, shortly petiolate, rigid, coriaceous, slightly
discolorous, sub-lustrous, green, becoming blue-
green with age, 5-9 cm long, 0.7-1.1 cm wide; petioles
0.2-1.0 cm long. Intermediate leaves lanceolate,
concolorous, lustrous, slightly blue-green, slightly

Muelleria

Rule

broader than the juvenile leaves. Adult leaves narrowly
lanceolate or lanceolate, sometimes falcate, semi-
erect, acuminate, uncinate, coriaceous (0.5-0.65 mm
thick), erect along the axis, sub-lustrous olive-green
or with a bluish tinge, 5-10 cm long, 0.8-1.5 cm wide;
new season's growth tips sub-glaucous; petioles terete,
0.9-1.4 cm long; venation moderately acute (35''-45°),
moderately reticulate; intramarginal vein c 1 mm
from the margin; oil glands large, numerous, irregular,
island. Inflorescences simple, axillary, 7-n-flowered,
along the main axis or along terminal, leafy branchlets;
peduncles angular or terete, slender, 0.6-1.1 cm
long. Floral buds ovoid to slightly clavate, petiolate,
lightly pruinose at anthesis, unscarred (petaline and
sepaline opercula intact), 4-6 mm long, c.3 mm wide;
pedicels 2-4 mm long, operculum conical, c. 2/3 the

length of and as wide as the hypanthium; locules 3 or
4; ovules in 4 vertical rows; filaments white; stamens
irregularly inflexed, all fertile; anthers sub-basifixed,
adnate, globoid, dehiscing through lateral pores.
Fruits hemisperical to cupular, rarely barrel-shaped,
pedicellate, 4-6 mm long, 3-4 mm diam; pedicels
1-3 mm long; disc descending; valves 3 or 4, enclosed.
Fertile seeds ovoid, slightly flattened, finely reticulate,
dark brown; hilum ventral (Fig. 5).

Flowering Period: Autumn.

Distribution and Habitat: Eucalyptus yarriambiack is
known from a single location between Brim and Beulah
in the southern part of the Victorian Mallee region
(Fig. 1). Its occurrence is on well-drained mallee loams
close to the Yarriambiack Creek which, due to low
rainfall of the area, is usually a dry water course.

Figure 5. Eucalyptus yarriambiack (a) tree; (b) bark; (c) seedling; (d) buds; (e) fruits

Vol 30(2) 2012

Five new Victorian eucalypts

Additional specimens examined: VICTORIA: NNE of Brim,
Wardles Road East, c. 200 m E of the railway line, K. Rule 1108,
l.vii.2008 (MEL); NW of Brim, Wardles Road West, c 1.1 km
W of Henty Hwy, K. Rule 1208, l.vii.2008 (MEL); NNW of Brim,
Starrocks Road, c. 600 m W of the Henty Hwy, K. Rule 1308,
l.vii.2008 (MEL); Henty Hwy N of Brim, c. 0.8 km S of creek
crossing, K. Rule 1408, l.vii.2008 (MEL); Henty Hwy, N of Brim,
c.2.7 km S of creek crossing, K.Ru/e 1608, l.vii.2008 (MEL).

Associated Species: Eucalyptus yarriambiack occurs in
pure stands but has contact with £ dumosa A.Cunn. ex
J.OxIey and £. largiflorens F. Muell. on the boundaries of
its distribution.

Etymology: The epithet, which is Aboriginal in
origin, is used as a noun in apposition and refers to
the Yarriambiack Creek close to where the new species
occurs.

Conservation Status: The total known number of
plants of £ yarriambiack is about 150, most of which
are mature trees. Seedling recruitment is minimal and
seedlings are absent along the stretch of the Henty
Highway where several of the mature trees occur. As far
as is known, all plants occur along roadside reserves and
thus are exposed to a variety of threats. In accordance
with lUCN criteria (lUCN 2001) the species is regarded as
'critically endangered'(CR).

Discussion: In habit and bark £ yarriambiack
resembles £ largiflorens, black box, which occurs nearby
in heavy soils along the Yarriambiack Creek. However,
the two differ markedly in a wide range of features,
particularly with the latter having duller, pendulous

foliage, terminal, compound inflorescences and scarred
buds.

Eucalyptus yarriambiack, whose features include adult
leaves with moderately reticulate venation and large,
irregular island glands, simple, axillary inflorescences
and 3-4-loculed fruits, clearly place it with the mallee-
boxes, a large group of taxa within the Series Buxeales.
The combination of features which distinguish the new
species from its relatives are its robust, often spreading
habit, its persistent box bark, its relatively narrow,
greenish juvenile leaves, its sub-lustrous, blue-green or
olive-green adult leaves, its sub-glaucous new growth
during the growing season, its lightly pruinose buds and
its small, pedicellate fruits.

Of the mallee-boxes, £ wimmerensis, which occurs
approximately 50 km to the west of £ yarriambiack, is
considered a close relative. Both have similar juvenile
leaves in shape and size and similar adult leaves in
shape, size and colour, as well as in venation and oil
gland patterns. However, £ wimmerensis differs by its
true mallee habit, its reduced amount of box bark or
being completely smooth-barked and its generally
larger fruits (Table 4).

The tree-like habit and persistent box bark also
suggest £ yarriambiack has an affinity with £ odorata.
However, that species differs by its broader leaves at all
stages, its non-pruinose, larger buds and its larger fruits.
Recent surveys of mallee-boxes in the Wimmera have
revealed that small, rough-barked mallees, previously
included within £ wimmerensis, are now considered

Key to the Brown Stringybarks

1 Operculum smooth or slightly scurfy (non-warty); locules 3 or 4.2

1: Operculum warty; locules 4-7.3

2 Adult leaves lanceolate, to 13 cm long; fruits 8-11 mm diam.£ arenacea

2: Adult leaves elliptical to ovate, to 8 cm long; fruits 5-8 mm diam.£ aurifodina

3 Bark stringy, extending to the secondary branches; buds shortly pedicellate.£ baxteri

3: Bark on trunk smooth or flaky (box-type); buds sessile.4

4 Bark flaky, box-like; adult leaves broadly lanceolate; fruits 11-13 mm diam. E. victoriana

4: Bark mostly smooth; adult leaves ovate to orbicular; fruits 11 -25 mm diam. 5

5 Disc steeply ascending; fruits 11-16 mm diam. E,serraensis

5: Disc more or less level with the rim; fruits 16-25 mm diam.£ verrucata

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to be forms of £ polybractea. These and other forms of
E. polybractea differ from £. yarriambiack by their mallee
habit and their strongly pruinose seedlings.

Series Pachyphloiae (Brown Stringybark
Complex)

Until 1988 the brown stringybarkcomplex had consisted
of only two recognised taxa; the widespread, variable £
baxteri (Benth.) Maiden & Blakely ex J.M.BIack, which
occurs from Kangaroo Island to the south coast of New

South Wales, and the similarly variable £ alpina Lindley,
which was regarded as being restricted to ridge-tops in
the Grampians Ranges. In 1988 £ arenacea Marginson
& Ladiges was segregated from £ baxteri to cater for
populations of robust, spreading mallees occurring on
inland sandy sites of western Victoria and adjacent areas
of South Australia. With regard to £ alpina, studies by
Marginson (1984), Ladiges and Humphries (1986) and
Marginson and Ladiges (1988) indicated that it consisted
of three species rather than one. Marginson (1984)

Table 4. Comparisons between E. yarriambiack and related species.

Characters

E. wimmerensis

E. yarriambiack

E.odorata

E. polybractea

Habit

mallee to 8 m tall

single or few- stemmed
tree to 10 m tall

mallee or small tree to

15 mtall

mallee to 10 mtall

Bark

smooth or often with
box-type stocking

box-type to major
branches

smooth or a box-type
stocking to various
heights

Juvenile leaves

Size

5-8 cm long, 0.8-1.6
cm wide

5-9 cm long, 0.7-1.1
cm wide

5-8-cm long 1-3 cm
wide

4-10 cm long, 0.8-1.5
cm wide

Colour, lustre

dull, blue-green or
sub-glaucous

sub-lustrous, green or
blue-green

dull, blue-green or
sub-glaucous

dull, blue-green or
glaucous

Petiole length

0.6-1.2 cm

0.2-1 cm

0.4-1.2 cm

0.2-1.3 cm

Adult leaves

Size

5-11cm long, 0.8-1.5
cm wide

5-10 cm long, 0.8-1.5
wide

6-12 cm long, 0.9-2
cm wide

5-9 cm long, 0.9-1.7
cm wide

Colour, lustre

sub-lustrous or
lustrous, green, blue-
green or olive-green

sub-lustrous, blue-
green or olive-green

dull or sub-lustrous,
green or olive green

dull, glaucous or sub-
glaucous

Intramarginal vein
(distance from margin)

c, 1 mm

c. 1 mm

1-2 mm

cl mm

Petiole length

0.8-1.5 cm

0.9-1.4 cm

0.6-1.5 cm

Colour of growth tips
during growing season

dull, blue green

sub-glaucous

blue-green

glaucous

Buds

Pruinosity

absent or light in one
southern form

light

absent

strongly present

Shape

ovoid to slightly
clavate

ovoid to clavate,

ovoid to slightly
clavate

Size

4-6 mm long, 3-4 mm
diam

4-6 mm long, c.3 mm
diam

5-8 mm long, 3-5 mm
diam

4-7 mm long, 3-5 mm
diam

Pedicel length

2-4 mm

2-5 mm

Fruits

Shape

cupular or sub-
cylindrical

hemispherical to
cupular, rarely barrel¬
shaped

cupular or barrel¬
shaped

cupular, sub-cylindrical
or barrel-shaped

Size

6-8 mm long, 4-6 mm
diam

4-6 mm long, 3-4 mm
diam

6-9 mm long, 5-7 mm
diam

4-7 mm long, 3-5 mm
diam

Pedicel length

1-3 mm

1-4 mm

100

Vot 30(2) 2012

Five new Victorian eucalypts

asserted that the materials used for the type specimen
of E. alpina, collected from the summit of Mt William,
were from a hybrid between £ baxteri and an unnamed,
related taxon and that the name £ alpino could no longer
be sustained. Chemical studies by Ladiges and Whiffin
(1993) confirmed the findings of previous researchers
and £ serraensis Ladiges & Whiffin. £ verrucata Ladiges
& Whiffin (referred to as £ verrucosa in the paper) and
£ victoriana Ladiges & Whiffin were subsequently
erected in 1993. The earlier name £ verrucosa Colla
rendered £ verrucosa Ladiges & Whiffin illegitimate and
was replaced by £ verrucata Ladiges & Whiffin in 1995.
In a concluding comment on £ baxteri, Whiffin and
Ladiges (1995) were of the opinion that £ baxteri held
no other segregate worthy of taxonomic recognition.

The distinguishing features upon which £ arenacea
was segregated from £ baxteri included its juvenile
leaves becoming glabrous later in seedling development
(hispid leaves persisting to at least 13 pairs in the former
and to 8 pairs in the latter), its leaves at all stages being
narrower, with the adult leaves more tapering to the
apex, its flower buds being non-warty with longer,
narrower pedicels, its peduncles also being longer and
narrower and its fruits being generally smaller, with a
less raised disc.

Further research into the variable nature of £ baxteri
has identified a new taxon occurring in the Victorian
Goldfields, a region whose populations were not
included in any of the studies of previous researchers
(cited above). A treatment of this new taxon is provided
below.

Eucalypus aurifodina Rule sp. nov.

Eucalypto arenaceae affinis foliis juveniUbus minoribus,
foliis intermediis ovatis vel sub-orbicularibus minoribus,
foliis aduftis ovatis vel elHpticis minoribus, alabastris
ovoideis minoribus, fructibus minoribus differt et
Eucalypto baxteri affinis foliis juvenilibus et adultis
minoribus, pedunculis angustoribus longioribus, alabastris
minoribus, operculis obtuso-conicis nomverrucatis,
fructibus pedicellatis minoribus differt.

Type: Victoria: Maldon Historical Reserve, c 200 m N
of Smiths ReefTrack along Tatt Town Track, 37°01'04"S,

144° 06' 02" E. K. Rule 3905 & £ Perkins 30.iv.2005. HOLO:
MEL

Habit small, single or multi-stemmed trees to 12 m
tall. Bark grey, stringy, sometimes latticed, extending to
secondary branches; inner bark reddish-brown. Seedling
leaves ovate, sessile, discolorous, lustrous and green
above, whitish below; hispid (hairs arising from raised oil
glands); margins hispid; seedling stems hispid. Juvenile
leaves ovate to broadly ovate, symmetrical, apiculate,
opposite for a few pairs, petiolate, hispid for less than
the 10”^ pair then becoming glabrous, discolorous,
lustrous and green above, pale green below, 3-5.5
cm long, 2-4 cm wide; petioles 0.4-1.1 cm long.
Intermediate leaves broadly elliptical to broadly ovate,
often basally oblique, slightly discolorous, lustrous and
green, regularly persisting in the mature canopy.'Adu/f
leaves elliptical or ovate, rarely ovate-lanceolate, often
basally oblique, apiculate, coriaceous, moderately
reticulate, lustrous and green, 4-8 cm long, 1.5-3
wide; petioles 1.2-2 cm long; lateral veins moderately
acute, 35®-40® to the mid-vein; intramarginal vein c.

2 mm from the margin; oil glands numerous, regular,
island. Inflorescences simple, axillary, 7-11-flowered;
peduncles slender, terete or slightly angular, 5-12 mm
long, contracting and thickening when bearing mature
fruits. Floral buds ovoid or slightly clavate, pedicellate,
unscarred (only a single operculum present), 5-7 mm
long, 2.5-3 mm wide; pedicels 3-5 mm long; operculum
conical (obtuse to moderately acute), smooth or slightly
scurfy (non-warty), flush with the hypanthium at the
abscission zone, 2.5-3.5 mm long, 2-3 mm wide; locules

3 or 4; ovules in 2 vertical rows; stamens inflexed, all
fertile; filaments white; anthers sub-basifixed, versatile,
reniform, dehiscing through oblique, confluent slits.
Fruits hemispherical, shortly pedicellate, 5-7 mm long,
5-7(-8) mm wide; pedicels 1-3 mm long; disc broad,
1.5-2 mm wide, slightly ascending, rarely flat; valves
3 or 4, triangular slightly exserted or level with the disc.
Fertile seeds black, irregularly sub-pyramidal, finely
pitted; hilum terminal (Fig. 6).

Flowering Period: Early autumn.

Distribution and Habitat Eucalyptus aurifodina
occurs in dry woodland communities in north-central
Victoria between Castlemaine and Avoca in areas once
exploited for their gold deposits (Fig. 1). Its preferred
habitat features gravelly soils on dry, stony slopes and
rises.

Muelleria

101

Rule

Figure 6. Eucalyptus aurifodina (a) tree; (b) adult leaves and buds; (c) seedling; (d) fruits; (e) buds.

Additional specimens examined: VICTORIA; Dunach Nature
Reserve, E of the Ballarat-Maryborough Road, K. Rule 15107,

2. xi.2007 (MEL); SW ofTalbot, c. 700 m NW along Norburys Road
from Lexton-Talbot Road, K. Rule 15207, 2.xi.2007 (MEL); 2.4 km
NE of Lexton on Lexton-Talbot Road, K. Rule 15307, 2.xi.2007
(MEL); 1.8 km WNW of Lexton on Lexton-Ararat Road, K. Rule
15407, 2.xi.2007 (MEL); 100 m N of LillicurWest Road-Sunraysia
Hwy intersection, K. Rule 15507, 2.xi.2007 (MEL); Muckleford
Flora Reserve, 31.viii.l981, A.C Beauglehole ACB68968 &
E. Perkins (MEL); Porcupine Ridge Road, K. Rule 4105 & E. Perkins,

3. V.2005 (MEL).

Associated Species: Eucalyptus microcarpa, E. nortonii,
E.macrorhyncha,E.tricarpa{lA.SJoUnsor\)lA.SJohnson
& K.D.Hill subsp. tricarpa, £ melliodora, £ dives,
E. polyanthemos (subsp. vestita and subsp. marginalis
Rule) and E.leucoxyfon subsp,pruinosa (F. Muell. ex Miq.)
Boland have been observed occurring with or adjacent

to the new species. Hybrids with £ macrorhyncha occur
to the north-east of Lexton.

Etymology.Jhe name is derived from the Latin aurifer
'gold-bearing' in reference to the new taxon being
located in the goldfields of north-central Victoria.

Conservation Status: In the western part of the
distribution the species has been collected from only
small remnant populations whose numbers appear
to have suffered from clearing for agriculture. A small
population occurs in the Dunach Flora Reserve, which
is administered by Parks Victoria. A second small
population occurs in the Lillicur State Forest whilst
the other western populations occur on roadside
easements or in small pockets scattered on private
land. On the other hand, the entire eastern part of the
distribution occurs in reserves under the management

102

Vol 30(2) 2012

Five new Victorian eucalypts

of Parks Victoria, namely the Smiths Reef Forest {Maldon
Historical Reserve), the Muckleford Nature Conservation
Reserve and the Castlemaine Diggings Nature
Conservation Park. The Smiths Reef Forest population is
by far the largest of the species, with numbers estimated
at more than 200 trees.The total number of mature trees
of the species in the known populations is estimated at
between 800 and 1000.

Given that its numbers are sparsely scattered and
probably amount to no more than a thousand plants,
in accordance to lUCN criteria (lUCN 2001), a status of
'vulnerable'(VU) is recommended.

Discussion: Eucalyptus ourifodina is distinguished by
its dry, gravelly, inland habitat, its habit of a small, single
or multi-stemmed tree, its relatively small leaves at all
stages, its slender, terete or slightly angular peduncles
to 12 mm long, its pedicellate, small, ovoid buds with
a smooth or slightly scurfy (non-warty), obtuse-conical
operculum and its relatively small, hemispherical fruits
with a moderately elevated disc and 3 or 4 valves.
It is considered to have some features intermediate
between £ orenacea and £ baxteri {Table 5).

The relatively long, slender peduncle, the non-
warty, thinly pedicellate buds and 3-4-loculed fruits of

Table 5, Comparisons between £ ourifodina and related species.

Characters

£ arenacea

E. baxteri

£ ourifodina

Habitat

sub-coastal and inland sandy
soils

coastal and sub-coastal
heavy soils

inland gravelly soils

Habit

usually robust spreading
mallees to 8 m tall

multi-stemmed coastal
shrubs to trees to 40 m tall

small, often multi-stemmed
trees to 12 m tall

Juvenile leaves

Shape

broadly ovate

broadly elliptical to broadly
ovate

broadly ovate

Size

6-10 cm long, 2-4 cm wide

6-13 cm long, 3-9 cm wide

3-5.5 cm long, 2-4 cm wide

Transition to glabrous leaves

by 14th node

by 8th node

by 10th node

Adult leaves

Shape

lanceolate or ovate-lanceolate

broadly ovate or ovate-
lanceolate

elliptical or ovate

Size

7-13 cm long, 2-3.5 cm wide

7-15 cm long, 2-5 cm wide

4-8 cm long, 1.5-3 cm wide

Peduncles

Length

6-12 mm long

2-4 mm long

5-12 mm long

Thickness

1.5-2.5 mm thick

2-3.5 mm thick

1.5-2 mm thick

Buds

Shape

clavate or slightly ovoid

clavate

ovoid or slightly clavate

Size

7-10 mm long, 3-4mm diam

5-8 mm long, 4-5 mm diam

to 5-7 mm long, c. 3 mm
diam

Pedicel length

3-5 mm long

0-4 mm long

3-4 mm long

Operculum shape

hemispherical or obtuse conical

hemispherical or obtuse-
conical

obtuse-conical, rarely conical

Operculum surface

non-warty

warty

non-warty

Fruits

Shape

hemispherical

hemispherical or globose-
truncate

hemispherical

Size

7-9 mm long, 9-11 mm diam

7-15 mm long, 8-15 mm
diam

5-7 mm long, 5-8 mm diam

Disc orientation

level or slightly ascending

slightly to prominently
ascending

slightly ascending

No of valves

3 or 4

4or5

3 or 4

Muelleria

103

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E. aurifodina suggest a close relationship with
E arenaceo. However, that species differs from the new
taxon by its sandy habitat, its delayed transition from
hispid to glabrous leaves during the seedling stage, its
longer, narrower (lanceolate), tapering adult leaves, its
larger buds and its larger fruits.

The new taxon shares some features with E baxteri,
namely the similar shaped pre-adult leaves and a similar
period of transition from hispid to glabrous leaves
during the seedling stage. However, E baxteri differs
by its wetter habitat (coastal and sub-coastal), its larger
leaves at all stages, its thicker peduncles, its warty buds,
which are often constricted at the abscission zone and its
larger fruits that are either sessile or shortly pedicellate
and have 4 or 5 valves.

Acknowledgements

A number of MEL staff contributed to the preparation
of this paper. Neville Walsh is thanked for his continued
support for this project and for providing the Latin
diagnoses. Val Stajsic, the late John Reid, and Jeff
Jeanes assisted with fieldwork and helpful discussion,
Catherine Gallagher facilitated access to MEL collections
and Alison Vaughan provided the distribution map. Both
Chris Jenek and Dermot Malloy of the Royal Botanic
Gardens Melbourne nursery are also thanked for their
work with the seedling trials. Special thanks are given to
Ern Perkins for his assistance in locating populations of
E aurifodina and E. conferta in the Castlemaine area and
Bill Molyneux of Dixons Creek for cultivating some forms
of E baxteri. Lastly, thanks are given to David Cameron
of the Arthur Rylah Institute for his assistance in the
field and for his advice regarding the recommended
conservation statuses for the treated taxa.

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Ladiges, P.Y. and Whiffin, T. (1995). A new name for Eucalyptus
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Vol 30(2) 2012

Five new Victorian eucalypts

Nicolle, D. (2000). New taxa of Eucalyptus, informal subgenus
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5,695-700

Muelleria

105

A revision of eastern Australian Bossiaea
(Fabaceae: Bossiaeeae)

Ian R. Thompson

The University of Melbourne, Parkville, Victoria 3010, Australia; email: i.thompson@unimelb.edu.au

(Host institution for duration of project: National Herbarium of Victoria, Royal Botanic Gardens Melbourne, BirdwoodAve,

South Yarra, Victoria 3141, Australia)

Abstract

Introduction

A revision of eastern Australian
Bossiaea (Fabaceae: Bossiaeeae) is
presented. Five new species, Bossiaea
alpina, B. dasycarpa, B. obovata,

B. peninsularis. and B. sericea, are
described, B. rhombifoUa subsp.
concoloris raised to the rank of
species as B. concolor, and B. cinerea
var. rigida is resurrected and raised to
the rank of species as B. tasmanica.
Bossiaea cordifolio, B. decumbens,

B. distichodada, and B. nummularia
are resurrected as species. An
informal infrageneric classification
and keys to groups and species are
presented, as well as illustrations,
images, and distribution maps.

Keywords: morphoiogy, taxonomy,
biodiversity, flowering plants, peas,
legumes.

Muelleria 30(2): 106-174 (2012)

Ku>mI

llotanic

Oardens

Mellnuirnc

Tribe Bossiaeeae (Fabaceae) is endemic in Australia and comprises
seven genera and 101 species. The tribe has a widespread distribution
in Australia, with most species occupying temperate and subtropical
latitudes. Morphological features defining the tribe include: stamens all
fused into an adaxially open sheath, anthers all dorsifixed and uniform
in size, and seeds with a laterally connected and distinctively lobed aril.
Standard and wing petals are fundamentally yellow, but commonly also
have reddish markings.

Bossiaea Vent, is morphologically diverse and is by far the largest genus
in tribe Bossiaeeae. It comprises 78 species (following the revision herein),
with centres of diversity in south-eastern Australia and south-western
Western Australia. Eastern species, with theexceptionofthe desert-dwelling
species B. walked, generally occur in regions with annual rainfall greater
than about 500 mm and mostly form part of the shrubby understorey of
open forests and woodlands. Platylobium Sm., which is endemic to south¬
eastern Australia, is clearly the closest relative of Bossiaea. The two genera
share the following distinguishing features: unifoliolate leaves (when
leaves are present), scales (fused stipule-pairs), terminal inflorescences
that are mostly 1- or 2-flowered and mostly on contracted axes (and so
appearing to be axillary), and brown, chartaceous bracts and bracteoles
which resemble scales. Bossiaea can be distinguished from Platylobium
by several features, including the less strongly compressed pods, the
lack of or less conspicuous wing development on the upper margin of
pods, and the stipules not both broad and deflexed. In addition, Bossiaea
generally has smaller leaves, fewer inflorescence scales, upper calyx-lobes
that are less conspicuously broader and longer than lower lobes, and
bracteoles that are generally more proximally inserted on the pedicel.
Within tribe Bossiaeeae, the lack of leaf development and the compressed
and sometimes winged branchlets seen in some species of Bossiaea are
unique. The leafy, eastern Australian species of Bossiaea have narrower
leaves than those of Platylobium, their inflorescences are subtended by
fewer inflorescence scales, and they tend to have smaller flowers.

The relationships within Bossiaea, and whether it constitutes a
monophyletic group separate from Platylobium is unresolved. The

106

Vol 30(2) 2012

Eastern Bossiaea

relatedness of eastern and western species is also
unclear; however, in the course of this study, preliminary
comparisons of the morphology of the eastern and
western species suggest that Bossiaea had already
diversified to a degree prior to the two regions becoming
botanically isolated. In Western Australia, there is
perhaps a greater amount of diversity than in the east,
and this is reflected by the fact that two genera, Lologe
Lindl. and Scottia R.Br., were erected in the early 1800s
to accommodate some species. Bentham subsequently
transferred these species to Bossiaea (Bentham 1864).
The Western Australian species were recently revised
(Ross 2006) and this resulted in an increase in the
number of species of Bossiaea In Western Australia
from 24 to 38. Bossiaea bossiaeoides (Benth.) A.B. Court
occurs in northern Western Australia and was described
in Ross (2006), but also extends across the Northern
Territory and into far north-western Qld. This species is
not included in the current revision.

The research described herein is the final taxonomic
instalment in a study encompassing all genera in tribe
Bossiaeeae. Revisions of Goodia Salisb. (Thompson
2011a), Platylobium (Thompson 2011 b), Muefleranthus

Hutch., Paragood/ol.Thomps.,Aen/cfop/?yfonA.T.Lee and
Ptychosema Benth. (Thompson 2011c) have recently
been published.

Taxonomic history (see Table 1): The genus Bossiaea
was erected in 1800 by the Frenchman Etienne Ventenat
when he described the eastern Australian species
R heterophylla Vent. From that time through until the
late 1880s, new taxa were named regularly, with Baron
Ferdinand Mueller and George Bentham being the main
contributors, Bentham in Flora Australiensis (Bentham
1864) named five new species, but also reduced to
synonymy four previously described taxa, and reduced
B. rosmarinifolia Lindl. to varietal rank. During this period,
several taxa now assigned to either Templetonia R.Br. or
Cristonia J.H.Ross in tribe Brongniartieae were initially
placed in Bossiaea. In the first three decades of the 20'^
century three new varieties were recognised, but then
there was a 53 year hiatus before the next new taxon,
B. oligosperma A.T.Lee, was described (Lee 1981). In 1991
Bossiaea arenicola J.H.Ross was described (Ross 1991),
and very recently there has been a burst of taxonomic
activity, with five new leafless species described (Ross
2008; McDougall 2009).

Table 1. Chronology of publication of new taxa in eastern Australian Bossiaea prior to the current revision.
Taxa that are treated as synonyms in the current revision, or that are unplaced, are presented in shaded boxes

(See also Index of scientific names).

heterophylla ClSOO)
obcordata^ {] 803)
scolopendrio' 808)
cinerea 08M)
prostrata {}8M)

foliosa m25)
buxifolia (\825)
ensata 0825)
lenticularis 0S25)
rhombifolia 0^25)
cordifolia^]827)
rosmarinifolia^ (1838)
nummularia 0S^9)

corinalis (1848)
distichoclada^ (1855)
cofd/gera (1856)
decumbens^^858)

walked OQbl)
brownii O^bA)
rupicola 0^64)
riparia (1864)
bracteosa 0^04)
kiamensis (1864)
neo-anglica 0^65)
arm/f//(1879)

scortechinii (1883)
stephensonii 0B87)

var. rigidcd (1903)
var. concolor^ (1908)

microphylfum' (1805)
lanceolawm' i\808)
ovafum'(1808)
cocc/ne(3(1813)

rotundifolia (1825)
linnaeoides 0B32)
tenuicaulis 0840)

humilis f1844)

fienderson/V (1866)

p/umoso (1893)

1800-1820

1821-1840

1841-1860

1861-1880

1881-1900

1901-1920

oligosperma 0980
arenicola 0990

vombata (2008)
bombayensis (2009)
fragrans (2009)
milesiae (2009)
gray! (2009)

var. stenoctadaf' (1928)

1921-1940

1941-1960

1961-1980

1981-2000

2001-2010

1. Originally placed in Platylobium.

2. Reduced to synonymy in Flora Australiensis (Bentham 1864).

3. Reduced In status In Flora Australiensis (Bentham 1864).

4. First described as a variety of B. cinerea, then treated as synonymous with 6 . obcordata.

5. First described as a variety of B. rhombifolia. Subsequently raised to B. rhombifolia subsp. concolor (Lee 1970).

6. Variety of B. heterophylla.

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Methods

This morphological study was based for the most part
on examination of herbarium material with the aid of
a dissecting microscope. In addition, trips into the field
were undertaken in Victoria and south-eastern New
South Wales to examine populations and to collect and
examine fresh material. Flowers of some herbarium
specimens were reconstituted using hot soapy water.
Specimens from AD, BRI, CANB, HO, MEL, NE, and NSW
were examined. All images presented in figures were
taken by the author using a Canon PowerShot A2000 IS
digital camera.

Type specimens cited have been seen by the author
unless indicated with n.v. The author has not examined
overseas material for purposes of typification except
via images, mostly digital and accessed on-line, and
further investigation is desirable in some instances. The
majority of images examined have been from K, but
images of material from G, BM and W have also been
examined. Illustrations have been designated as the
holotype of a species in several instances. Searches of
herbaria in the future may reveal specimens associated
with these illustrations, but there has been no indication
of their existence in documents I have read.

Taxa with a current conservation classification are
indicated in the Distribution and habitat sections
below, and new taxa that are likely to warrant such a
classification are also indicated here.

Explanatory notes for keys and descriptions:
Measurements in keys and descriptions are based on
pressed herbarium material and, unless otherwise
specified, are based on branchlets (see Notes on
morphology) and the structures borne on them. For
leafless species, cladode-scale measurements refer to the
longest scales of a branchlet, which occur in the middle
third. Descriptions of stipules, bracts and bracteoles refer
to their abaxial (outer) surface and width measurements
of bracts and bracteoles were taken of the structures
in their natural shape rather than flattened out.
Descriptions of stipules refer to their appearance soon
after maturity and before they become degraded with
age. Also in the descriptions, the hairiness of stipules,
scales, as well as bracts, bracteoles and calyces refer to
their abaxial (outer) surface, not including the margins.
The descriptions of convexity of bracts and bracteoles is
also based on the abaxial view. The length of the calyx
does not include the length of the receptacle (see Figure
1e). Descriptions of petal colour and marking pattern
are partly based on field examination and examination
of photographs and partly on interpretation of pressed
specimens. The colour of wings and keel is that of their
abaxial (outer) surface. Descriptions of petal colour
and seed morphology are based on a limited survey of
specimens for many species. The shape of pods given
in descriptions is the two dimensional shape seen in
profile.

Table 2. A new, informal classification of eastern Australian Bossiaea.

Group A

Group B

Group C

Group D

Foliosa subgroup

1. B. foliosa

2. B. distichodada

3. B. sericea

4. S. afpina

Onerea subgroup

5. B. cinerea

6. B. rosmarinifolia

7. B. cordifolio

Kiamensis subgroup

8.6. kiamensis

Cordigera subgroup

9. B. lenticuloris

W. B. cordigera

Buxifolia subgroup

/ 1. B. decumbens

12. B. buxifolia

13. B. neoanglico

Prostrata subgroup

14. B. nummularia

15. B. prostrata

Scortechinii subgroup

16. B. dasycarpa

17. B. scortechinii

18. B. obovata

Obcordata subgroup

19. B.tasmanica

20. B. obcordata

Group E

Group F (Leafless)

Stephensonii subgroup

21. B.stephensonii

Heterophylla subgroup

22. B. heterophylla

23. B. rhombifolia

24. B. concolor

Brownii subgroup

25. B. carinalis

26. B, rupicola

27. B. brownii

28. B. oligosperma

Arenicola subgroup

29. B. arenicola

Ensata subgroup

30. B. ensata

31. B.scolopendria

32. B. peninsularis

33. B. ormitii

34. B. riparia

Fragrans subgroup

35. B. fragrans

36. B. milesiae

Bracteosa subgroup

37. B. bombayensis

38. B. grayi

39. B. vombata

40. B. bracteosa

Walkeri subgroup

41. B. waikeri

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Vol 30(2) 2012

Eastern Bossiaeo

The term l:w ratio refers to the length of a structure
divided by its width and is here expressed as a quotient,
e.g., I:w ratio 3 indicates that the structure is 3 times
longer than wide, and I:w ratio 5-8 indicates that the
structure ranges from being five times longer than wide
to eight times longer than wide.

Results

The results of this morphological study are presented
in the taxonomic section below. In summary, the
pattern of morphological variation identified calls
for the recognition of eleven additional species,
including five new species, one taxon elevated in
rank, one taxon resurrected and elevated in rank,
and four species resurrected. An informal two-tiered
infrageneric classification has also been developed, and
is summarised in Table 2.

The most taxonomically important characters
identified in this study were: habit; branch architecture,
compression and spininess; indumentum density;
stipule dimensions, shape, texture and orientation;
presence or absence of leaves and scales; phyllotaxy;
leaflet size, shape and articulation; apiculum
morphology; inflorescence-axis length; pedicel length
and robustness; bracteole shape, persistence and
insertion position; calyx-lobe shapes and relative sizes;
petal lengths and coloration; pod length, thickness and
indumentum; pod-stipe length; and aril morphology.

Notes on morphological features

BRANCHLETS: Branchlets are here defined as the
branches bearing inflorescences in their axils.They vary
from terete to compressed.The branchlets and branches
of leafless species are mostly broadly winged and are
termed cladodes. Compressed branchlets generally
become more terete in subsequent years. Ridges or leaf
decurrencies are often evident on branchlets and when
well-developed are generally pale. In leafy species,
branchlets are sometimes moderately compressed but
there is usually little or no wing development. Bossiaea
stephensonii F. Muell. is an exception as it typically does
develop narrow wings. The wings are mostly much
wider than the terete or mildly compressed central
region. The pale margins of the wings of these cladodes
correspond to the decurrent ridges of leafy species.

In some species, branchlets are striate due to raised
parallel venation or develop a whitish epicuticular wax
as they mature. In leafless species in particular, this wax
layer can ultimately lift in fragments varying from small
flakes to large sheets, e.g., see Figure 12d.

Bossiaea kiamensis Benth. is unique amongst eastern
species in having minutely bulliform branchlets and
branches (Fig. 4e).

INDUMENTUM: Hairs are simple, basifixed, straight,
or less often curled, and 0.2 to 1 or rarely to 2 mm long.
They are mostly white, but occasionally pale yellow
or coppery. The inner surface of calyx-lobes has an
indumentum of short, curled hairs, and the margins
of scales, bracts, bracteoles and calyx-lobes are almost
always ciliolate. Hairs are generally similar throughout a
plant with the exception of pod-valves which often have
hairs longer than on other structures. The orientation of
hairs can vary within a species from antrorse-appressed
to spreading. Some species more commonly have
spreading hairs while others more commonly have
appressed hairs, but orientation is not generally a
reliable taxonomic character. The minute tubercles
evident on upper surfaces and margins of leaves of
several species are persistent hair-bases.

Taxa displaying a relatively distinctive indumentum
include; the four species in the Foliosa subgroup (Group
A), which have pale yellow or coppery hairs on ovaries
and pods; Rstep/ienson/V, which has relatively long hairs;
B. brownii Benth. which has long hairs emerging from a
layer of curled hairs; and 6. cordigera Benth. ex Hook f.
which has scattered short, curly hairs. Leafless species
are commonly glabrous at maturity, but scattered
hairs are generally present on developing cladodes,
at least on the margins. Some members of the Ensata
subgroup develop hairs on the apex of the keel-petals.
Leafy species that are often virtually glabrous include
B. lenticularis Sieber ex DC. and species in the
Heterophylla subgroup.

STIPULES AND SCALES; Stipule morphology is quite
variable. Most stipules have a ciliolate apex even if the
surfaces are glabrous. Stipules are generally persistent,
but generally darken and become degraded with age.
Stipules are inserted on the lateral expansion of the
branch on which a leaf is inserted. In all but one species
insertion of the pair of stipules is opposite; however, in
B. cordifolia they are inserted more adjacent to one
another (Fig. 4f). The stipules of this species are also

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o o

Figure 1, Aspects of morphology, a. Petiole and petiotule showing variation in their articulation, (i) articulation obscure; (ii)
as previous except for a spur, (iii) articulation marked by a change in diameter from petiole to petiolule, (iv) articulation geniculate,
(v) as previous but also with a ridge, (vi) as previous but with petiole spurred; b. Reproductive organs (example species Bossiaea
heterophylla). (i) calyx, cut and opened out, (ii) standard, (iii) wing, (iv) keel, (v) androecium, (vi) gynoecium, (vii) pod, (viii) cross-
section of pod. All approximately x2; c. Seeds (i) Aril-base broad (Cinerea subgroup), (ii) Small seed, aril-base narrow, strongly
curved lobe (Buxifolia subgroup), (iii) Aril knobbly and with lobe oblique, obscuring gap between lobe and base on one side
(a walken), (iv) Seed typical of several groups (drawn from B. heterophylla); d. Inflorescences (bracts and bracteoles not shown),
(i) sessile, with one pair of inflorescence scales, (ii) as previous but with shoot development below scales, (iii) as previous except
stipules and leaf developed instead of scales; (iv) as for (i) but with multiple pairs of inflorescence scales, (v) rudimentary on-
growing of axis adjacent to flower, (vi) 2-flowered cluster, (vii) 2-flowered raceme-like inflorescence with rudimentary growing-
on of axis, (viii) as previous but with on-growing axis leafy. Individual flowers could be interpreted as solitary and axillary in this
arrangement; e. Calyces (showing one upper lobe, one lower lateral lobe and lower median lobe (appearing as half actual width
due to the lateral view; base of calyx marked with a dashed line), (i) upper lobes triangular and the same size as lower lobes
(Bracteosa subgroup), (ii) upper lobes triangular, broader than lower lobes {B. arenicola), (iii-vii) upper lobes c. quadrate; (iii) lower
lobes relatively long and lobe-apices filiform (Scortechinii subgroup), (iv) upper lobes oblong, narrow (Foliosa subgroup), (v)
upper lobes moderately broad, apex nearly truncate (eg., Heterophylla subgroup), (vi) upper lobes broad, broadening markedly
from base, with apex rounded (eg., Cinerea subgroup), (vii) as previous but upper lobes relatively long (eg., Cordigera subgroup).

Drawings not precisely to scale, but all within the range x4 to x5.

Vol 30(2) 2012

Eastern Bossiaea

unusual in the degree of deflexion. The angle the stipule
makes with the petiole varies within a small range in most
species, but this measure can be useful taxonomically
(e.g., see Figure lOh, i).The angle is difficult to assess if
stipules are markedly recurved or deflexed.

In species of the Brownii subgroup of Group F,
margins of stipules are clear and membranous and
become recurved to revolute and generally distorted.
In species of Groups B and C, stipules are commonly
very narrowly triangular, often filiform distaily, reddish,
and tend to become recurved. Bossiaea stephensonii
has exceptionally large, green, erect stipules. Stipules in
the Scortechinii subgroup of Group D are significantly
smaller than those of B. stephensonii, but are otherwise
similar in appearance.

Scales are formed by the fusion of stipules and are
present at nodes of the cladodes of leafless species and
on the inflorescence axes of all species. They are also
often present at the most basal nodes of branches of
leafy species. At nodes occupied by a scale a leaf is not
developed. The scales on cladodes of leafless species
become progressively longer with distance from the
base up to a certain point, and are longest in the middle
third of the cladode. The basal scales of a cladode are
sometimes clustered.

In leafy species there are two scales at the base of
inflorescences; they are small and are mostly shorter
than and often hairier than the adjacent bract. They are
often only partially fused and sometimes a minute leaf
rudiment is developed between the two scale halves to
produce a trifid arrangement. Because of the small size
and hairiness these inflorescence scales are difficult to
observe. Occasionally stipules and a small to normal¬
sized leaf are present on the axis instead of a scale. In
leafless species, there are often multiple, overlapping
pairs of inflorescence scales and the most distal ones are
relatively large.

PHYLLOTAXY: Phyllotaxy is mostly distichous in
eastern Australian Bossiaea. Most species have regularly
alternate leaves; however, three species have opposite
leaves, and the three species in the Cinerea subgroup
have variable phyllotaxy. Although leaves are inserted
in a spiral arrangement in the Cinerea subgroup, the
laminas tend to become oriented in one plane.

LEAVES: Leaves in eastern Australian Bossiaea are
unifoliolate, although in a small proportion of species
the articulation, /.e., the articulation between the

petiole and the petiolulate leaflet, is obscure and the
leaf appears to be simple (Fig. la, i). The articulation
is recognisable in most species by either a change of
angle (geniculate articulation; Fig. la, iv-vi), change in
diameter (a non-pulvinate petiole terminus connecting
to a pulvinate petiolule; Fig. la, iii), or by the presence
of a ridge or spur on the petiole (Fig. la, v & vi and
Fig. 4e). Species in the Cordigera subgroup have
relatively slender petioles.

The laminas of leaflets are generally small, but are
highly variable in shape. For branchlets, lengths range
from a minimum of 1 mm in B, alpina I. Thomps. and
B. distichoclada F. Muell. to a maximum of 30 mm in
B. rosmarinifolia. Bossiaea carinalis Benth. has the
broadest leaflets (to 13 mm). Species in the Buxifolia
and Brownii subgroups have leaflets with asymmetric
bases (Fig. lOe). The asymmetry is sometimes quite
subtle and is often variably evident within a plant.
Margins of leaflets are mostly flat or recurved, but are
sometimes revolute, e.g., in the Cinerea subgroup,
and rarely incurved to involute. In B. rhombifofia
DC. B. concolor (Marden & Betche) I. Thomps.
B. heterophylla Vent, and B. stephensonii, the margin
generally has a fine pale rim that is generally only
evident with microscopic examination. The apex of
leaves is highly variable in shape and in the degree of
development of an apiculum. The apiculum is either
straight or downcurved. The stoutest and longest
apicula are seen in the Cinerea subgroup and these are
commonly pungent (Fig. 4).

Minute tubercles evident on the upper surface and
margins of some species are hair-bases. Scattered
pale dots, presumably glands, are sometimes evident
on the upper surface of leaves of pressed specimens
(when viewed under magnification); however, different
pressing methods appear to influence the visibility of
these dots. This gland-dotted appearance is perhaps
most consistently seen in species in Group D.

INFLORESCENCE ARCHITECTURE (Fig. 1 d):
Inflorescences typically appear to be axillary; however,
they are interpreted here as terminal inflorescences
arising from contracted axes.The evidence for this axis is
commonly no more than the presence of a pair of scales
below the pedicel of a flower.

In a number of leafless species, a few to several
overlapping pairs of inflorescence scales are developed,
while in several species, e.g., species in the Cordigera

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Thompson

subgroup, the scales are separated from the axil by
an axis up to c. 3 mm long. Less commonly, a short
inflorescence axis is more obviously developed, with
several, often leafy, nodes present. In most species,
only a single inflorescence develops from an axil, but
occasionally, and especially in the Bracteosa subgroup,
multiple inflorescences arise from an axil.

Inflorescences are mostly single-flowered, with two-
flowered clusters occurring occasionally in several
species. However, in species in the Scortechinii subgroup
a raceme-like arrangement of two or three flowers can
develop, with the inflorescence axis continuing beyond
the distalmost flower. This axis may be rudimentary at
flowering or it may have developed into a leafy axis.
Occasionally in this subgroup a solitary, truly axillary
flower, subtended by a leaf or scale, may develop along
an otherwise leafy shoot (as in Fig. Id, viii but with only
one flower).

BRACTS AND BRACTEOLES: A single bract is present
at the base of each pedicel and commonly is partly
obscured by one of the subtending inflorescence
scales. The two bracteoles are inserted on the pedicel at
different points depending on the species, ranging from
basal to almost at the summit. Bracts and bracteoles
are commonly slightly to strongly convex abaxially and
they always have ciliolate margins. They are glabrous
adaxially (on the inner surface) but are often hairy at
first abaxially. Bracteoles are commonly opposite, but
sometimes one is inserted up to c. 1 mm distal to the
other. This sub-opposite arrangement is quite often
seen in B. carinalis. In the Buxifolia subgroup bracteoles
tend to be both inserted towards the upper side of the
pedicel rather than on opposite sides, and they are often
more widely divergent (Fig. 6d). In B. ensata Sieber ex
DC. and B. scolopendria (Andrews) Sm. bracteoles are
somewhat fleshy. In B. arenicola the two bracteoles fuse
to form a single structure (Fig. lOn). In B. distichoclada
bracteoles are relatively large and a colourless, recurved
margin is sometimes evident.

In most leafy species bracteoles are persistent until
well after anthesis and they often persist through to
mature fruit stage. In contrast, seven of the 12 leafless
species have caducous bracteoles. Because these
species with caducous bracteoles have multiple pairs of
inflorescence scales and because they are structurally
very similar to bracts and bracteoles, it can be difficult
to determine whether bracts and bracteoles have been
shed. Sometimes, although abscissed, they remain

lodged in position. In B. milesioe KL. McDougall and
B. walked abscission scars are usually visible on the
pedicel, but in other species these scars are concealed
by scales.

RECEPTACLE AND CALYX (Fig. le): The receptacle
is the dilated distal end of the pedicel and is a well-
developed obconical structure 0.5-1 mm long (labelled
in Figure le). The junction between receptacle and
calyx is generally identifiable in pressed specimens.The
calyx-tube is 1-4 mm long and is variously shorter than,
equal to, or longer than the lobes. In a few species, e.g.,
B. heterophyllo and 6 . cadnalis, the calyx is often slightly
glaucous and in others, notably B. heterophyllo, B. ensata
and B. scolopendria, the calyx-tube commonly has
broad, longitudinal red stripes aligned with the inter¬
lobe sinuses.

Upper calyx-lobes are highly variable in eastern
Australian Bossiaea. Except for four species with
triangular lobes, the upper calyx-lobes are somewhat
four-angled but with considerable variation in shape,
mostly relating to broadening from the base and the
degree of expansion beyond the lateral angle. Major
variants are illustrated in Figure le. In the final two
examples (Fig. le vi & vii) there is expansion of the lobe
beyond the lateral angle. When this occurs the apex is
always broadly rounded.The calyx shown in Figure le
vii resembles the morphology of Platylobium. Lower
calyx-lobes are triangular to narrow-triangular and are
relatively uniform in morphology. In several species,
notably B. scortechinii F. Muell. and B. dasycarpa CT.
White ex I.Thomps. the apices of both lower and upper
lobes have a filiform extension. In B. scortechinii, the
lower median lobe is often distinctly longer than the
lower lateral lobes. The calyx-lobes of the Bracteosa
subgroup are distinctive in being all triangular and of
similar size and shape as well as being largely brown
and chartaceous.

COROLLA; Outlines of petal shapes of B, heterophyllo
are shown in Figure 1b and are fairly typical of the
shapes in the majority of species. The standard-limb is
generally slightly broader than long except in Group A
species in which it is more or less circular.The unopened
standard in Group A species is also folded so that the
lateral margins merely touch rather than overlap as is
typical of other species.

Two fairly common patterns of coloration of petals are
shown in Figures 101 {B, oligospermaPJ. Lee living) and 8d

112

Vol 30(2) 2012

Eastern Bossiaea

{B. prostrata R.Br. pressed). In the former, the standard is
yellow adaxially (internally) except for a red flare around
the throat. Wings are flushed reddish or brownish
abaxially over much of their length, and the keel is a
darker purple-red distally. On the standard a red band
sometimes runs vertically through the throat to divide
it in two. The throat also commonly has red flecks at the
base. The abaxial (outer) surface of the standard mostly
has some degree of pink to red colouring. Sometimes,
as is shown in Figure 8d, pale lines corresponding to
the course of veins radiate from the flare and interrupt
an otherwise red surface. Wings are sometimes entirely
yellow except for some pink markings towards the base.
Five species, B. arenicofa and the four species in Group A,
always have entirely yellow petals, while three species
in the Scortechinii subgroup, especially B. scortechinii,
are typically yellow or with relatively little red marking.
Yellow-petalled mutant plants are occasionally recorded
for species that normally have red markings.

PODS: The upper margin of pods is variable in
thickness and in the degree of development of vertical
ridges. Sometimes the ridge is restricted to the suture
line only, and there may be a sulcus formed each
side of this ridge. If the ridge is much higher than
wide it approaches the dimensions of a wing, as the
ridge is generally referred to in Platylobium. Pods of
B. carinalis could almost be described as having wings
(Fig. 10k). Pods with thickened valves and broadened
upper margins are only seen in Group E and in a few
species in Group F. In most groups the upper margin is
0.5-1 mm wide, whereas it ranges from 1 to 3 mm wide
in species in Group E. Extremes in the range of widths of
the upper margin are shown in Figure lOg with a pod of
6. rhombifolio placed beside a pod of R buxifolia.

The outer surface of valves commonly has slightly
raised transverse venation evident with magnification;
however, in species in Group B the venation is usually
indistinct. The inner surface of pod valves is mostly
smooth and glabrous; however, in several species in
Group E spongiose tissue forms between valves creating
a partition between the seeds (Fig. lOf).

There appears to be some variation in the degree of
revolute rolling of valves post-dehiscence. The rolling
appears to gradually develop post-dehiscence. In some
species the valves persist on the plant post-fruiting and
are present in the next flowering period as cylinders
with the exposed inner surface being silvery.

SEEDS (Figs 1 c & 4g-i): Seeds are relatively uniform in
shape and they range in length from 2 to 6 mm. Mature
seeds are brown to blackish and are commonly mottled
(Figs 4g-i, 10c). Seeds become considerably shorter but
plumper just prior to maturity. When examining seeds
of herbarium records it may be difficult to tell if that final
change of shape had occurred. Some measurements of
seed length may turn out to be excessively long for this
reason.The aril is also fairly uniform in shape and relative
size. There is some variation in the length of its base and
the degree of overhang and curvature of the lobe. The
oblique arching or asymmetry of the recurved margins
of the lobe, which is a normal feature, is evident in Figure
4h. The aril of B. walkeri is unusual in being slightly
knobbly and with the gap between lobe and base being
hidden when viewed from one side.

Taxonomy

In the descriptions below, species are ordered according
to morphological similarity and, to further emphasise
points of similarity, they have also been placed in six
informal groups and 16 subgroups (Table 2).The groups
are in some instances somewhat weakly defined,
whereas the subgroups are well-defined and likely
to reflect close relationships between members. The
epithet of the most familiar or most widespread species
in a subgroup is adopted for the name of the subgroup,
eg.,The Prostrata subgroup is named after R prostrafa.

Bossiaea Vent., Descr. PI. Nouv. 1:7(1800)

Type: Bossiaea heterophylla Vent.

Bossieua, orth, var. Pers.

Boissiaea, orth. var. Lem.

Scottia R.Br., in W.T.Aiton, Hortus Kew., edn 2, 4: 268
(1812). Type: 5. dentata R.Br. = R dentata (R.Br.) Benth.

Lalage Lindl., in J.Lindley, Edwards's Bot. Reg. 20:1.1722
(1834). Type: L ornato Lindl. = B. ornata (Lindl.) Benth.

[All taxa historically placed in either Scoff/a or Lalage
are endemic to Western Australia.]

A circumscription of Eastern Australian species

Subshrubs, shrubs or small trees, sometimes leafless,
sometimes rhizomatous. Indumentum commonly
developed but variably persistent on branchlets and
leaves, sometimes developed on pedicels and ovaries

Muelleria

113

Thompson

Key to informal groups of eastern Australian Bossiaea

1 Plants leafless (or occasionally with a few leaves developed from lower nodes of broadly winged stems).Group F

1: Plants leafy........ 2

2 Leaves all regularly opposite, with nodes well-spaced.3

2: Leaves all regularly alternate or arranged irregularly.4

3 Leaflets with l:w ratio s 2.Group B

3: Leaflets with l:w ratio 0.8-1.2.Group C

4 Standard petal to c. 8 mm long, all yellow; upper calyx-lobes < 1 mm wide; pods < 10 mm long, hairy all over;

ovaries and pods with hairs commonly pale yellow or coppery (mostly tablelands to alps).Group A

4: Standard petal > 8 mm long and/or with red markings, or if ever not with either feature then upper
calyx-lobes > 1 mm wide; pods > 10 mm long, hairy all over or more often with valve faces glabrous;
ovaries and pods with hairs white.......5

5 Apex of leaflets narrowly acute with a robust, sometimes pungent apiculum; bracteoles to c. 1 mm long,

not or hardly longer than broad, inserted more than halfway along pedicel.Group B

5: Apex of leaflets not entirely as above or if ever nearly so then branchlets compressed; bracteoles mostly

> 1 mm long, longer than broad, inserted variously..6

6 At least some branchlets spinescent or subspinescent (tapering to a blunt point).Group D

6: Branchlets not spinescent or tapering.7

7 Lower calyx-lobes at least as long as the calyx-tube AND the upper lobes; calyx hairy.Group D

7: Lower calyx-lobes shorter than calyx-tube and/or shorter than the upper lobes; calyx glabrous or hairy.8

8 Ovaries and pods hairy at least on margins; prostrate or weakly erect shrubs to c. 0.5 m high (higher if supported);

pods thin, with upper margin < 1 mm wide; pod-stipe generally < 3 mm long.-.Group C

8: Ovaries and pods glabrous, or if ever hairy then tall shrubs with flowers entirely yellow and bracteoles
fused into a single structure; generally erect shrubs, mostly > 0.5 m high; pods slightly to very thick,
with upper margin > 1 mm wide; pod-stipe generally > 3 mm long.....Group E

and abaxial surfaces of stipules, scales, bracts, bracteoles
and calyces, rarely developed on the apex of keel-petals;
margins of structures such as stipules, scales, bracts,
bracteoles and calyx-lobes almost always ciliolate;
hairs simple, mostly straight but occasionally curled or
crumpled, white or rarely yellow or coppery. Branchlets
terete or com pressed, sometimes with leaf decurrencies,
sometimes broadly winged (and then called cladodes),
rarely spinescent, sometimes developing epicuticular
wax.Sf/pu/eserect,reflexedorrarelydeflexed,herbaceous
or scarious, fused to form scales below inflorescences
(inflorescence scales) and, in leafless species, at all nodes
along cladodes. Leaves mostly distichous, alternate or
less often opposite or irregularly arranged, unifoliolate
but with the articulation sometimes obscure, or leaves
absent. Inflorescences mostly appearing axillary, but

interpreted as terminal on a contracted or very short or
rarely more elongate axis bearing 2 or less often 4-10
scales; inflorescences mostly 1-flowered, occasionally
2-flowered, and very occasionally with flowers in a
raceme-like arrangement; bract and bracteoles mostly
scarious; bract at base of pedicel, slightly shorter than
bracteoles; bracteoles variously inserted, persistent or
caducous; receptacle obconical, generally distinct. Calyx
with tube shorter than to longer than lobes; upper lobes
partly fused, mostly broader and often longer than the
triangular lower lobes; petals clawed; standard and
wings yellow, commonly with reddish or purple-brown
markings; standard with limb oblate or occasionally
c. circular; standard mostly equal to or slightly longer
than wings; keel equal to, slightly longer, or occasionally
much longer than wings, pale or more often reddish

114

Vol 30(2) 2012

Eastern Bossiaea

Key to Group A

1 Leaflet halves markedly convex, with margins commonly partly revolute, with upper surface generally with

scattered minute tubercles all over; lower surface never covered by closely appressed hairs.2

1: Leaflet halves flat or only slightly convex, with margins not revolute, with upper surface lacking tubercles

or tubercles near-marginally only; lower surface commonly with a close-appressed to sericeous indumentum.3

2 Stipules < 1.5 mm long; leaves commonly broad-ovate; bracteoles s 1 mm long, not exceeding receptacle. ^.B.foliosa

2: Stipules > 1.5 mm long; leaves commonly transversely oblong-elliptic; bracteoles > 2 mm long,

extending well onto the calyx.2, B. distichodada

3 Lower surface of leaflets largely to totally covered by a closely appressed, generally sericeous indumentum;
bracteoles usually with some hairs, s 1.5 mm long except in Brindabella Ranges and Coolamon Plain

(Australian Capital Territory and New South Wales) where sometimes up to 2.5 mm long. 3,B,sericea

3: Lower surface of leaflets glabrous or with scattered hairs near midline only; bracteoles glabrous,

^ 2 mm long (Victoria only).4. B. aipina

especially distally; stamens fused to form an adaxially
open sheath; anthers all dorsifixed and ±uniform in size;
ovary 2-20-ovulate; style slender, upcurved; stigma
small. Pods stipitate; body compressed, with valves
and margins variously thickened, glabrous, hairy on
margins, or hairy all over, sometimes with spongiose
tissue partitioning seeds internally; upper margin often
slightly to strongly ridged. Seeds plump, ellipsoid or
slightly reniform, with a conspicuous aril; aril with a lobe
arising at one end and arching over the aril-base.

Group A

Shrubs, mostly erect but sometimes nearly prostrate;
branchlets terete, without decurrent ridges, commonly
moderately hairy. Stipules often striate. Leaves with
lamina small, generally not longer than broad, markedly
discolorous, often lustrous or sublustrous above.
Inflorescences: axes contracted, with scales 2; pedicels
short; bracteoles persistent, inserted proximally,
sometimes relatively large. Flowers relatively numerous,
relatively small; upper calyx-lobes narrowly oblong, not
expanded beyond lateral angle, with lateral angle acute;
petals entirely yellow; anthers small {0.2-0.3 mm long).
Pods short-stipitate; body circular to broad-elliptic, hairy
all over, with hairs commonly pale yellow to coppery.
Seeds 1 or 2 per pod (Fig. 2).

Group A is a well-defined and distinctive group of four
species. It is also identified here as the Foliosa subgroup
to emphasise the close relationship between members.
It occurs in south-eastern New South Wales and eastern
Victoria at moderate to high altitudes, and contains

the only species to extend into the alpine zone (Fig. 3).
It is essentially synonymous with B. foliosa A. Cunn. as
historically circumscribed, Le., in Flora Australiensis and
in subsequent state floras. Descriptions in recent state
floras erroneously state that 6 . foliosa, Le„ in the historic
sense, has caducous bracteoles.

1. Bossiaea foliosa A.Cunn., in B.Field, Geogr.
Memoirs New South Wales: 347 (1825)

Type: not designated. [Protologue: 'Brushy forest¬
land near Bathurst'.] New South Wales. Near Bathurst,
A.Cunningham, 1822; lectotype (here selected):
K 000278308, image seen in Kew Herbarium Catalogue;
probable isolectotypes: K 000278306, K 000278307,
K 000278309, images seen in Kew Herbarium Catalogue,
NSW 566395.

Erect shrubs to c. 1.5(-2) m high, with inflorescences
borne typically on a ±regular series of short side-
branchlets; branchlets erecto-patent to spreading,
c. 0.4 mm wide, with a moderately dense indumentum
of straight or wavy hairs c. 0.2 mm long; epicuticular wax
not or hardly developed. Stipules triangular to narrow-
triangular, 0.4-1 (-1.5) mm long, erect, brown, often
darker medially, hairy, glabrescent, 3 or more-nerved,
but venation often obscure; stipule-petiole angle
60-90°. Leaves: petiole 0.3-0.7 mm long; articulation
not or slightly geniculate, not ridged; lamina commonly
broad-ovate, less often c. orbicular, squarish or
transverse-oblong, 1.5-3 mm long, 1.5-3.5 mm
wide, with l:w ratio mostly 0.8-1.1, with each lamina

Muelleria

115

Thompson

Figure 2. Group A. a. Bossiaea distichodado {J.H.Ross3631 MEL); b. B. distkhodada, pod {RJ.FIetcher376 MEL); c. B. sericea
{R.O.Makinson 920 MEL); d. B. alpina, flowers and new growth {RKBorley. 19.xi.l980 MEL); e. & f.adaxial and abaxial views of leaves.
Top row: R foliosa {I.Crawford 769 MEL), and R distichodado UH.Ross 3632 MEL); bottom row: R sericea {R.O.Makinson 920 MEL)
and R alpina {D.E.Afbrecht 5195 MEL); g. R foliosa, flowers {KJ.Fitzgerold 75 MEL). Scale bars: a, c = 10 mm, b, d-g. = 2 mm.

half strongly convex, markedly discolorous; base
symmetrical, truncate to cordate; margin often revolute
each side of midline in distal two-thirds, otherwise
recurved; apex broadly rounded or truncate: apiculum
hardly developed, generally pointing down; upper
surface minutely tuberculate, sometimes wrinkled
(which may obscure tubercles), with venation not
evident, with gland-dotting not evident, soon
glabrescent; lower surface minutely white-dotted, with
scattered hairs on midrib and extending laterally onto
lamina, but often only sparsely hairy or glabrous towards
periphery; hairs often slightly wavy, loosely appressed
to somewhat spreading. Inflorescences: axes contracted;
bract c. 0.5 mm long, c 0.5 mm wide, strongly convex;
pedicel 0.5-2.5 mm long, glabrous or occasionally with
scattered hairs; bracteoles persistent, broad-ovate,
0.5-1 mm long, with l:w ratio c 1. divergent, inserted at
base, strongly convex, with venation obscure, glabrous
or sparsely hairy distally, red-brown. Calyx 2-3.5 mm
long, hairy, with tube c. equal to lobes; upper lobes

0.8-1.8 mm long, c. 0.8 mm wide; sinus 0.3-1.2 mm
deep; lower lobes 0.6-1 mm long, c. 0.5 mm wide, flat;
petals all similar in length, all entirely yellow; standard to
c. 6 mm long; wings c. 1.5 mm wide; keel c. 2 mm wide;
ovary hairy, 2-ovulate. Pods: stipe c. 1.5 mm long; body
c. circular, 5-8 mm long, 5-7 mm wide, with scattered
pale or light golden hairs c 0.6 mm long on valves and
margins; upper margin c. 0.5 mm wide, with ridge to
c. 0.3 mm high. Seeds 2.0-2.8 mm long, 1.5-1.8 mm
wide; aril 1 -1.2 mm long, c. 0.8 mm high, with base c. 0.6
mm long, with lobe curving 60-120° (Fig. 2e-g).

Selected specimens from c. 60 examined: NEW SOUTH
WALES: The Mullions Range, 10 km NE of Mullion Creek RS
(c 24 km NNE of Orange), R.Coveny 10234, 14.X.1978 (NSW);
29.2 km from Yass Rd along Nottingham Rd toward Tumut,
HJhompson 906 & P.OIIerenshaw, 27.i.1987 (CANB, MEL, NSW);
Monaro Hwy between Nimmitabel and Bombala, 2 km S from
turn-off to Snowy Mtns Hwy, G.Stewart 416, 4.xii.1984 (CANB,
MEL); Tantangara turn-off, c. 16 km S of Kiandra, BMeiner
494, 20.xii.1960 (CANB); Dry Plains Rd, c. 25 km from Cooma,
R.W.Purdie 5623, 23.xi.2002 (CANB); Hume and Hovell Walking

116

Vol 30(2) 2012

Eastern Bossiaea

Track, Burrinjuck Nature Reserve, KJ.Fitzgerald 75, 8.xi.1997
(CANB, HO, MEL, NSW). AUSTRALIAN CAPITAL TERRITORY:
Old Boboyan Rd (south), 1.2 km W of turn-off from Canberra-
Adaminaby road, M.D.Cnsp 9285 & LG.Cook, 4.xiL2000 (CANB,
NSW); between Canberra and Lake George, E.Gauba, 30.X.1949
(CANB). VICTORIA: Craigie Bog Road S of New South Wales
border, A.CBeauglehole 34822, 23j<i.l970 (MEL); Limestone
Creek, J.Stirling, 1.xii.l882 (MEL); source of Mitta Mitta River,
J.Stirling,}8B2 (AD).

Flowering period: Flowers from Octoberto December.

Distribution and habitat: Occurs in the central and
southern tablelands of New South Wales and in far
eastern Victoria at altitudes of about 800-1200 metres
a.s.l. (Fig. 3a). Grows in often stony soils in open forest
and woodland.

Notes: Bossiaea foliosa is distinguishable from other
species of the group by a combination of leaflet-shape,
texture of the upper surface and indumentum of the
lowersurfaceof leaflets, small bracteoles with somewhat
rounded apices, hairy calyces, and glabrous pedicels.
The leaflets are commonly ovate because margins
are revolute distolaterally. The leaflets of members of
Group A are compared in Figure 2e-f. The bract and
bracteoles of B. foliosa are inserted close together
and are divergent; this creates a cupular arrangement
from which the pedicel emerges and generally clearly
exceeds. The bract is often trifid, an uncommon shape
for bracts but commonly seen in inflorescence scales.
Bossiaea foliosa and B. distichodada have leaflets with
a pale lower surface. Under magnification, this pallor is
seen to be due to minute white rings closely crowded
over the surface.

Hybridisation: A specimen collected from Haydons
Bog near Delegate, in far eastern Victoria {Bauerlen, 1899
NSW) may be a hybrid between B. sericea I.Thomps, and
B. foliosa A. Cunn.

Typificot/on: There are two sheets at K with probable
type material of B. foliosa. Although the collector's
name is not specified for some pieces, I consider that all
pieces were collected by Cunningham on his expedition
from Port Jackson to Bathurst in 1822-1823, and were
probably all from a single gathering near Bathurst.
On one sheet a piece coded as K 278308 is labelled
'Brushy forest land near Bathurst' and so matches the
description in the protologue. A.B. Court annotated
this piece as 'the type' in 1967, but Lee (1970) while
discussing 'the Holotype' did not clearly indicate

whether she was referring to this piece. All pieces are of
similar diagnostic value, but I here formally designate K
278308 as the lectotype of B. foliosa based on the close
correspondence between the label and the protologue.
The contents of a small envelope on the sheet near the
lectotype have not been seen. On the same sheet, the
piece in the top right hand corner K 278307 is labelled
as a Cunningham collection, Cunningham 730/1822,
while for the two pieces of K 278309, the collector is
not indicated. Both are probable isolectotypes. The
second sheet at K bearing probable type material,
coded as K 278306, has three pieces and is a duplicate
of K 278307 based on similarities in the labelling, and
so also probably an isolectotype. At the time of writing,
the catalogue had incorrectly identified the collection
number for this specimen as Cunningham 136. The
number 130 is written on the label but a comma from
the line above has caused it to appear like 136.

2. Bossiaea distichodada F.Muell., Trans. Phil.
Soc. Victoria 1:39 (1855)

Type: not designated. [Protologue: 'In the Australian
Alps from the Mitta Mitta to the tributaries of the Snowy
River'.] Victoria. Bogong Mountains, F.Mueller, date
unknown (probably 1854); lectotype (here selected):
MEL 20321.

Residual syntypes: Victoria. Mitta Mitta, F.Mueller,
i.l854: MEL 20323, MEL 20326, MEL 20327 (all
B. distichodada): Victoria. Upper Avon, Gippsland,
F.Mueller, xi.l854, MEL 20320 (mixed sheet of
6. distichodada and B. a/p/no); Victoria/New South Wales.
Snowy Mountains, F.Mueller, possibly i.1855: MEL 20322
(mixed sheet of B. distichodada and B. alpina; locality
given is possibly an error); Victoria. Locality unknown,
F.Mueller. MEL 20324; Victoria. Mt Wellington, F.Mueller,
xi.1854: MEL 20325 (mixed sheet of B. sericea and
B. alpina).

Bossiaea foliosa sensu G.Bentham, FI. Austral. 2: 160
(1864), and subsequent Australian authors, pro parte,
non sensu stricto.

Erect shrubs to c. 1 m high, with inflorescences borne
typically on a ±regular series of short side-branchlets;
branchlets sub-erect to erecto-patent, c. 0.6 mm
wide, with a dense indumentum of straightish hairs
c. 0.3 mm long; epicuticular wax not developed. Stipules

Muelleria

117

Thompson

Figure 3. Distributions of species in Group A. a. Bossiaea foliosa; b. B. distichoclada; c. B. sericeo; d. B. alpina.

triangular, 1.5-4 mm long, erect, divergent, sometimes
slightly reflexed, brown, glabrous, 5-10-nerved, but
venation generally obscure; stipule-petiole angle
variable, difficult to assess. Leaves: petiole 0.2-0.5 mm
long; articulation obscure; lamina obcordate, reniform,
or c. circular, 1 -2.5 mm long, 2-4 mm wide, with l:w ratio
mostly 0.6-0.9, with each lamina half strongly convex,
markedly discolorous; base symmetrical, broadly
rounded to cordate; margin recurved proximally,
generally revolute distally; apex broadly rounded,
truncate or emarginate; apiculum not or hardly
developed; upper surface minutely knobbly and often
wrinkled, with venation obscure, with gland-dotting
not evident; lower surface glabrous, or sometimes with
hairs on midrib, often white-dotted. Inflorescences: axes
contracted; bract 1.5-3 mm long, 1 mm wide, strongly
convex abaxially; pedicel 1 -2 mm long, hairy; bracteoles
persistent, narrow-elliptic, (2-)2.5-4.5 mm long, with
l:w ratio 2-3, loosely appressed to calyx, inserted near
base, strongly convex, with venation obscure, hairy
medially, brown. Calyx 2.5-3.5 mm long, hairy, with tube
slightly longer than lobes; upper lobes 1-1.5 mm long,
c. 0.8 mm wide; sinus c. 1 mm deep; lower lobes

1- 1.5 mm long, 0.5-0.7 mm wide, flat; petals all similar
in length, all entirely yellow; standard to c. 8 mm long;
wings 1.5-2 mm wide; keel 2-2.5 mm wide; ovary hairy,

2- ovulate. Pods: stipe 1 mm long; body c. circular or
broad-elliptic, 5-8 mm long, 5-6 mm wide, with rusty-
orange hairs c. 1 mm long on valves and margins; upper
margin c. 0.4 mm wide, ±unridged. Seeds c. 2.5 mm long,
1.5-2 mm wide; aril 1-1.2 mm long, c. 0.7 mm high,
with base 0.6-0.8 mm long, with lobe curving c. 90°
(Fig. 2a,b,e, f).

Selected specimens from c. 60 examined: NEW SOUTH
WALES: 9 km along Cascade Trail from the Alpine Way, 500
m S of Cascade Creek ford, AJ.Whalen 283, 14.xii.l998 (CANB,

DNA, MEL). VICTORIA: Blue Shirt Creek, 100 m downstream
of Nunniong Rd, 5 km WSW of Mt Nunniong, KMenkhorst s.n.,
12.i.1985 (MEL); c. 16 km E of Mt Hotham on road to Omeo,
J.Cullimore 246, 17.1.1968 (MEL); c. 9 km E of Little Mt Tambo,
c 3 km NE from junction of Currawong Rd and McDougall
Spur Track, F.B.Davies 609, 10.xi.1988 (AD, CANB, MEL, NSW);
Brumby Point, Nunniong Plateau, c. 80 km NNE of Bairnsdale,
J.H.Willis, 13.xi.l 964 (MEL); Forlorn Hope Track, S of Benambra-
Wulgulmerang road, A.C.Beaugleho!e 36194, 18.i.l971 (MEL);
Spring Hill Track, 12 km SSW of Mt Tambo, Splitters Range,
G.W.Carr 10247, 8.xii.l984 (MEL).

Flowering period: Flowers from Decemberto January.

Distribution and habitat: Occurs in eastern Victoria
from Mt Hotham east to The Cobberas and in far south>
eastern New South Wales where it occurs near Thredbo
in Kosciusko National Park (Fig. 3b). There Is a record of
B. distichoclada labelled Buffalo {Robbins MEL 521828)
which is part of a mixed collection with Rser/cea. There
is some doubt about the validity of the label. Grows in
montane to subalpine woodland.

Notes: Bracteoles of S. distichoclada are large and
have a conspicuous indumentum of appressed hairs
which is densest medially (Fig. 2a). Bracteoles extend
to the calyx-lobes or sometimes beyond this point.
Bracts and bracteoles tend to be more brittle than in
other species and often are split in pressed specimens.
Margins of stipules and bracteoles tend to fray with age.
The indumentum of pods (Fig. 2b) is more conspicuously
coppery than in other species. Leaves are closest in most
respects to those of R foliosa, but the margins are rolled
differently and do not create the ovate shape typically
seen in the latter. Specimens with very long stipules
and bracteoles have been recorded from the Nunniong
Plateau in eastern Victoria.

Hybridisation: Probable hybridisation has been
recorded between R distichoclada and R sericea in the

118

Vol 30(2) 2012

Eastern Bossioea

Bogong High Plains in north-eastern Victoria [IH.Ross
3635-7, 3640, all MEL) and further east at Limestone
Creek {N.G.Walsh 2867 CmS, MEL, NSW).

ryp/ficat/on; There are eight sheets at MEL, according
to Lee (1970), which Mueller may have used when
describing the species. Several of them have multiple
pieces and are mixed collections. The majority of pieces
conform to Mueller's description, but several pieces
do not, and are in fact specimens of 6. alpina, or in one
case, B. sericea, Lectotypification is therefore required for
B. distichoclada, and MEL 20321 is here selected. The
sheet contains a single piece with pods, and was labelled
as B. distichoclada Ferd Mueller by the author.

3. Bossiaea sericea I.Thomps., sp. nov.

A B. foiiosa A.Cunn. folioHs non ovatis laevibus superne
sericeis inferne, leguminibus majoribus, seminibus
majoribusdiffert.

Type: Victoria. 11.8 km by road from Rocky Valley
Dam wall towards Omeo, R.OMokinson 920, 3.xii.1991;
holotype: MEL 234474; isotype: BRI, CANB 9106346.

Bossiaea foiiosa sensu G.Bentham, FI. Austral. 2; 160
(1864), and subsequent Australian authors, pro parte,
non sensu stricto.

Erect shrubs to c. 2 m high, with inflorescences borne
typically on a ±regular series of short side-branchlets;
branchlets erecto-patent, c. 0.5 mm wide, with a
dense indumentum of straightish hairs c. 0.3 mm long;
epicuticular wax not developed. Stipules triangular,
0.7-2 mm long, erect, brown, glabrous, 4-10-nerved;
stipule-petiole angle 30-60® Leaves: petiole 0,5-1.5
mm long; articulation slightly to strongly geniculate,
not ridged; lamina c. circular, oblate, obcordate, broadly
obovate, or broadly quadrangular, (1.5-)2-4 mm long,
1.5-5 mm wide, with l:w ratio mostly 0.8-1.0, flat to
moderately folded, with each lamina half flat to slightly
convex, markedly discolorous; base symmetrical,
truncate to slightly cordate; margin flat to moderately
recurved, smooth or tuberculate, with a pale rim; apex
broadly rounded, truncate or emarginate; apiculum
minute, generally pointing slightly down; upper surface
smooth throughout or tuberculate at margins (visible
in abaxial and sometimes adaxial view), with venation
obscure, with gland-dotting not evident, glabrous;
lower surface evenly sericeous, sometimes densely, or

with indumentum sparser near margins; indumentum
usually somewhat persistent. Inflorescences: axes
contracted; bract 0.7-2.2 mm long, 0.5-1 mm wide,
strongly convex; pedicel 1-3.5 mm long, glabrous or
hairy; bracteoles persistent, ovate, narrow-ovate or
oblong, 0.7-2.5 mm long, with l:w ratio 1-3, mildly
divergent, inserted near base, strongly convex, with
venation mostly obscure, glabrous or with hairs towards
apex, brown. Calyx 2-3.5 mm long, hairy throughout or
glabrous except for lobes, with tube equal to or slightly
longer than lobes; upper lobes 0.9-1.5 mm long, 0.8
mm wide; sinus 0.5-1 mm deep; lower lobes 0.5-1
mm long, 0.6 mm wide, flat; petals all similar in length,
all entirely yellow (sometimes with pink tinges on
margins); standard to c. 8 mm long, with limb as long as
broad; wings 1.5-2 mm wide; keel 2-2.5 mm wide; ovary
hairy, 2- or 3-ovulate; style 2.5-4 mm long. Pods: stipe
1-2.5 mm long; body broad-elliptic, 6-10 mm long,
4-8mmwide,withrustyhairsoramixtureofpaleand rusty
hairs c. 1 mm long throughout; upper margin c. 0.5 mm
wide, with ridge to c. 0.2 mm high. Seeds often reniform,
(2.5-)3-4 mm long, c. 2 mm wide; aril 1.5-2 mm long,
1-1.2 mm high, with base 0.7-1.2 mm long, with lobe
curving 90-140° (Fig. 2c, e, f).

Selected specimens from c. 200 examined: AUSTRALIAN
CAPITAL TERRITORY: Mt Gingera, Brindabella Range, M.Evans
2565, 29.xi.1966 (AD,BRI, CANB, MEL, NSW); Mt Franklin, c. 0.5
km from chalet in direction of peak, T.R.LaHy & B.Lafoy 452,
23.xi.l 994 (CANB, NSW); 22 km S of Picadilly Junction, W.Bishop
584, 29J<ii.l987 (CANB, MEL, NSW); lower slope of Mt Ginini,
Brindabella Range, H.Coveny T1549 & P.Hind, 19.i.l983 (CANB,
MEL, NSW). NEW SOUTH WALES: Coolamon Plains, G.Singh,
29.xi.1979 (CANB); W side of Port Philip Fire Trail, 0.7 km from
Long Plain Rd, 3.1 km N of Rules Point, P.CJobson 5439 &
P.H.Weston,2^^.^99S (NSW); 16.7 km along Geehi Dam Rdfrom
the Alpine Way, Kosciusko National Park, RJohnstone 1523 &
AE.Orme, 20.i.2005 (NSW); 9 km along Cascade Trail from the
Alpine Way, Kosciusko National Park, AJ.Whalen 293, 14.xii.1998
(CANB, NSW); Maragle Range, Mt Black Jack, F.E.Davies 479
&S.Walton, 21.i.1988 (CANB, MEL, NSW); Constance's hut site,
Burrungubugge River, Kosciusko National Park, AMLyne 230,
28.i.1991 (CANB, MEL, NSW); near Eucumbene Lookout, Snowy
ms, R.A.Goode520, 17.xi.l961 (NSW); Mt Kosciusko, J.M.Curran,
i.1896 (NSW); near Tooma Pond, Kosciusko National Park,
AM.Ashby37l8 ,21 j<i.l 970 (AD). VICTORIA: Delegate River Fen,
near Old Bendoc-Bonang road, E.A.Chesterfield 42, 13.xi.1983
(CANB, MEL); Mt Bogong, G.Weindorfer, xii.1903 (MEL); Wall of
Death, Hotham Heights, D.E.AIbrecht 4948, 8.iv.l992 (CANB,

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119

Thompson

MEL); High Plains Rd, 0.8 km N of Falls Creek village, N.G.Walsh
3285, 5.ii.l992 (MEL); between Mt Anderson and Mt Pinnibar,
A.CBeauglehole 41569 & K.CBogers, 24.ii.1973 (MEL); Stoney
Creek Fire Trail, 4.5 km NNE of The Horn, Mt Buffalo, N.G.Walsh
3294, 18.ii.l992 (MEL); E side of The Horn Rd, 1.6 km N (by
road) of The Horn, Mt Buffalo National Park, P.CJobson 4032,
31 -xii.l995 (MEL, NSW); Mt Buffalo beside Lake Catani, M.AJodd
216, 26j<i.l974 (MEL); Camping Ground, Mt Buffalo, K.Czornij
387, 5.xii.l971 (AD); Mt Buffalo summit, CJ.Shepherd 203,
1.xii.l 965 (CANB).

F/oiven/igpenorf; Flowers from December to January.

Distribution and habitat: Occurs in north-eastern
and far eastern Victoria, far south-eastern New South
Wales, and along the western margin of the Australian
Capital Territory. In Victoria it extends from Mt Buffalo
ESE to the Delegate River east of Bonang, and in New
South Wales its range extends from the Kosciusko region
NNE to the Brindabella Ranges (Fig. 3c). Grows in heaths,
shrubland and woodland, often bordering grasslands.
Grows mostly above c. 800 m a.s.L and extends into the
alpine zone.

Etymology: The epithet refers to the indumentum
of the abaxial surface of the leaves (from Latin, sericeus,
silky).

Notes: Bosslaea sericea is moderately variable in leaf
shape and bracteole length, with three geographically
segregated forms. Forms from Mt Buffalo in north¬
eastern Victoria and the Brindabella Ranges and
adjacent Coolamon Plain in the A.C.T. and south-eastern
New South Wales have smaller, more angular leaflets
that are more tuberculate on margins and less densely
sericeous than the larger, more rounded leaflet form
from locations such as Mt Hotham, the Bogong High
Plains and ranges in far east Gippsland in Victoria, and
from Kosciusko National Park in New South Wales. The
leaflet-apex of the Brindabella and Mt Buffalo forms is
usually not emarginate, whereas the apex in the higher
altitude form can be. In far eastern Gippsland, the
rounded leaflet form generally has larger stipules than
other populations. The leaf representing B. sericea in
Figure 2e and f (bottom left) is of the form. The leaves
of the other two forms range between this shape and
the shapes shown for B. alpina (bottom right). The form
from the Brindabella Ranges and Coolamon Plain differs
from the other two by having relatively long bracteoles
(1.6-2.5 mm long compared to 0.7-1.5 mm long).

Leaflets of B. sericea lack tubercles (persistent hair-
bases) or the tubercles are only present near the margin.

The surface is commonly quite smooth but also can b^
slightly uneven, at least when dry, due to some faint
dark ridges between secondary veins.

Hybridisation: Probable hybridisation between
B. sericea and B. distichoclada q.v. has been recorder
in the Bogong High Plains in north-eastern Victoria
U.H.Ross 3635-7, 3640 all MEL) and further east at
Limestone Creek {N.G.Walsh 2867 CANB, MEL, NSW),
A specimen from Haydons Bog near Delegate {Bauerlet)^
1899 NSW) may be a hybrid between B. sericea anq
B. foliosa. A small sterile plant collected from an
unknown locality (Australia felix) in Victoria {F.Mueller
MEL 668111) is possibly a hybrid between Bossiaeq
sericea and B. prostrata.

4. Bossiaea alpina I.Thomps., sp- nov.

A B. foliosa A.Cunn. p/an f/s humiHoribus, foliolis laevibus
superne, calyce glabro, bracteolis longioribus differt.

Type: Victoria. Surveyors Creek Camp, D.E.AIbrecht
5195, 15.xii.l 992; holotype: MEL 2017313; isotype: CANB
n.v.

Bossiaea foliosa sensu G.Bentham, FI. Austral. 2: 16o
(1864), and subsequent Australian authors, pro parte^
non sensu stricto.

Diffuse shrubs to c. 0.5 m high, with inflorescences borne
typically on a ±regular series of short side-branchlets;
branchlets erecto-patent, c. 0.5 mm wide, with a
moderately dense indumentum of straightish hairs c.
0.3 mm long; epicuticular wax not developed. Stipules
triangular, 0.7-1.5 mm long, erect to divergent, brown,
glabrous, 5-10-nerved; stipule-petiole angle 30-60o.
Leaves: petiole 0,2-0.5 mm long; articulation slightly
geniculate, not ridged, sometimes obscure; lamina c.
square or transversely oblong to oblong-elliptic, 1-2
mm long, 1-2.5 mm wide, with l:w ratio mostly 0.8-1.0,
flat or more often concave, with each lamina half flat or
gently convex, markedly discolorous; base symmetrical,
c. truncate; margin slightly recurved, smooth or
minutely tuberculate; apex truncate or broadly
rounded, apiculum to c. 0.2 mm long, pointing forwards
or down; upper surface smooth, with venation obscure,
with gland-dotting not evident, glabrous; lower surface
glabrous or sparsely hairy and then glabrescent, without
white-dotting. Inflorescences: axes contracted; bract 2-3
mm long, 1.5-2 mm wide, moderately convex; pedicel

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Vol 30(2) 2012

Eastern Bossiaea

1.5-2.5 mm long, glabrous or with scattered hairs;
bracteoles persistent, narrow-ovate or narrow-oblong,
2-3.5 mm long, with l:w ratio 2-3, loosely appressed over
calyx, becoming slightly divergent at anthesis, inserted
near base, moderately convex, many-nerved, glabrous,
brown. Calyx 3-3.5 mm long, glabrous or sparsely hairy
near apex of lobes, with tube c. equal to lobes; upper
lobes 1.5-2 mm long, 0.8 mm wide; sinus c. 1-1,5 mm
deep; lower lobes c. 1.5 mm long, 0.8-1 mm wide, flat;
petals all similar in length, all entirely yellow; standard
to c. 8 mm long; wing 1.5-2 mm wide; keel 2-2.5 mm
wide; ovary hairy, 2-ovulate. Pods {only immature pods
seen): stipe c. 1 mm long; body c. circular, 6 mm long, 5
mm wide, with whitish hairs c. 0.5 mm long throughout;
upper margin c. 0,4 mm wide, with ridge not evident.
Seeds not seen (Fig. 2d-f).

Selected specimens from c. W examined: VICTORIA.
Tamboritha Saddle, near Chester's Hut, S of Bennison Plain,
N.GWalsh 974, 20.xi.l 980 (MEL, NSW); Echo Flat, Lake Mountain,
N.G.Walsh 908, 17.xii.l981 (MEL); The Bluff, c. 13 km SE of Mt
Buller, T.B.Muir 960, 28.xii.1959 (MEL); unnamed track 100 m
NE of Howitt Rd, 1.5 km NW of Guy's Hut, R.HSarley, 19.xi.l 980
(MEL); near Moroka Gap, 1.6 km SW of Mt Wellington, T.B.Muir
3744, 13.i.l965 (MEL); Lake Mountain, BJ.Corroll, 22.xii.1965
(CANB); Dry Creek, Howitt Plains, r.M.Whaite64, 8.1.1949 (NSW).

Flowering period: Flowers from Decemberto January.

Distribution and habitat Occurs in south-eastern
Victoria at Lake Mountain, Mt Buller and the Howitt
Plains area (Fig. 3d). May warrant recognition as a rare
species. Grows in subalpine or alpine heathland or
heathy woodland.

Etymology: The epithet refers to the occurrence of
this species in alpine and subalpine environments.

Notes: Bossiaea olpina is most closely related to
B. sericea but is distinguished from that species by its
smaller more sparsely hairy leaves, shorter petioles,
larger, more distinctly striate and glabrous bracteoles,
and glabrous calyx (Fig. 2d). Based on label data and
some field observations, B. alpina has a considerably
more diffuse habit than other members of Group A.

Group B

Erect shrubs; branchlets mostly terete, with decurrencies
not well-developed, moderately hairy, with hairs
straight. Stipules relatively narrow and often with apex
filiform, commonly recurved or deflexed, reddish-
brown, hairy abaxially. Leaves with phyllotaxy variable,
with articulation often spurred; lamina markedly
discolorous, with margins often recurved to revolute,
with midrib and apiculum generally robust, sometimes
pungent. Inflorescences: axes mostly contracted, with
scales 2; bracts and bracteoles small, nearly flat; pedicels
often long and slender; bracteoles persistent, 0.8-1.5
(-2) times longer than wide, inserted in middle or distal
thirds of pedicel. Calyx mostly glabrous; upper lobes
broadening markedly from the base, broader than long,
generally expanded beyond lateral angle; standard
generally 1-2 mm longer than wings and keel; wings
c. equal to keel; anthers relatively large. Pods with stipe
equal to or slightly longer than calyx; body generally
c. elliptic or oblong-elliptic, ±glabrous, with valves
smooth (transverse venation generally obscure). Seeds
mostly 2 or 3 per pod; aril long-based. (Fig. 4.)

Group B contains four species divided into two
subgroups. Members of the group are most readily

Key to Group B

1 Leaves all regularly opposite, with nodes well-spaced.

^: Phyllotaxy not as above, irregular and varying from alternate, to ±opposite to whorled on a single plant.

.8. B, kiamensis
.2

2 Leaflets with l;w ratio > 8, with upper surface smooth; leaflet-articulation marked by a spur and by

being geniculate (Grampians Ranges, Victoria only). B. rosmarinifolia

2: Leaflets with l;w ratio < 8, with upper surface usually minutely tuberculate; leaflet-articulation obscure

or marked by a spur but not geniculate....^

3 A small spur marking position of leaflet articulation in at least some leaves; leaflets with i:w ratio mostly

> 3; stipules inserted opposite each other; wings purplish-brown; pedicels commonly with some hairs.5. B. cinerea

3: Leaflet articulation obscure (spur not developed); leaflets with l:w ratio mostly < 3; stipules inserted
relatively close to each other (commonly forming a deflexed V); wings yellow except for reddish streaks
basally; pedicels usually glabrous.7.B. cordifolia

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121

Thompson

Figure 4. Group B. a. Bossiaea cinerea {T.B.Muir5545 MEL); b. B. cordifolia iT.B.Muir5091 MEL); c B. kiamensis {LR.Telford 237 CANB);

d. B. rosmarinifolia {T.B.Muir 866 MEL); e. 6. kiamensis, leaf and stipules {I.R.Telford 237 CANB); f. B. cordifolia, leaf and stipules
{H.Forde, x.1905 NSW); g & h. B. cordifolia, seed, left and right lateral views {I.RJhompson 1466 MEL); i. B. cinerea, seeds attached
inside pod {I.RJhompson 1436 MEL). Scale bars: a, b, d = 5 mm, c = 10 mm, e, f, i = 2 mm, g, h = 0.5 mm.

122

Vol 30(2) 2012

Eastern Bossiaeo

identified by leaf and bracteole morphology. Species
in Group B occur in south-eastern New South Wales,
southern Victoria, far south-eastern South Australia, and
Tasmania (Fig. 5).

The Cinerea subgroup (species 5-7) is well-defined.
The three members have irregular phyllotaxy, with the
arrangement varying from alternate to opposite to
whorled on a plant, and all have long, slender pedicels
and small, distally inserted bracteoles, Bossiaeakiomensis
(8) forms a subgroup on its own and is placed in Group
A because of similarities to the Cinerea subgroup in
leaf and pod morphology. In other respects, notably its
opposite leaves, it is closer to the Cordigera subgroup
of Group C.

The Cinerea subgroup

5. Bossiaea cinerea R.Br., in W.T.Aiton, Hortus
Kew., 2nd edn, 4:268 (1812)

Type: not designated. [Protologue: 'Native of Van
Diemen's Island, Robert Brown, Esq. Introd. 1805'.]
Tasmania. Port Dalrymple, R.Brown, 1.i.1804; lectotype
(here selected): BM 000885933, image seen in JSTOR
Plant Science.

Residual syntypes; Tasmania. Derwent River, R.Brown,
1802-05: BM 000885939, MEL 1528714, MEL 1528715,
MEL 1528716; possible residual syntype: Tasmania.
Locality unknown: CANB 00278253 (see discussion
below).

Bossiaea coccinea Bonpl., in A.Bonpland, Descr. PL
Malmaison 128, t. 52 (1813). Type: not designated.
[Protologue:‘Habitat in Nova Hollandia' Described from
a plant presumably cultivated at Jardin de la Malmaison,
Paris, France.] Holotype: t. 52 in Bonpland, Descr. PL
Malmaison 128 (1813).

Bossiaea tenuicauHs Graham, Edinburgh New Philos.
J. 29: 171 (1840); B. cinerea van tenuicauHs (Graham)
J.M.Black, FL S. Australia 2: 304 (1929). Type: not
designated. [Protologue: 'This plant was raised at the
Botanic Garden, Edinburgh, from Van Diemen's Land
seeds sent by Mr Cooper, Wentworth House, in Apr.
1836'.]

Erect shrubs to c. 2 m high, with inflorescences borne
typically on longer branchlets rather than a regular
series of short side-branchlets; branchlets erecto-
patent, c. terete or angular, 0.5-1 mm wide, with hairs

0.3-0.8 mm long; epicuticular wax generally absent.
Stipules narrow-triangular to filiform, 1-3 mm long,
erect or more often becoming markedly recurved,
reddish, hairy, with venation obscure; stipule-petiole
angle mostly 30-60°. Leaves alternate, sub-opposite,
opposite or in whorls of 3 in varying proportions on a
single plant; petiole 0.2-0.5 mm long; articulation not
geniculate, obscure except when marked by a spur
0.1-1 mm long; spur present on most leaves, or rarely
uncommon on a plant; lamina narrow-ovate to narrow-
lanceolate or narrow-triangular, 10-20 mm long, 1.5-6
mm wide, with l:w ratio mostly 3-8, slightly convex
each side of midrib, becoming strongly convex laterally,
markedly discolorous; base symmetrical, - broadly
rounded or truncate; margin recurved or revolute,
occasionally undulate, sometimes with a few persistent
hairs; apex narrowly acute; apiculum 0.4-1.2(-2) mm
long, sometimes pungent, sometimes somewhat brittle^
pointing forward or slightly dov^n; upper surface smooth
or tuberculate, with venation commonly raised, with
gland-dotting not evident, glabrescent; lower surface
usually hairy. Inflorescences: axes contracted or rarely
to c. 1 mm long; bract c. 1 mm long, c. 0.5 mm wide,
slightly convex; pedicel 2-11 mm long, mostly sparsely
hairy; bracteoles persistent, mostly broad-ovate, 0.2-1
mm long, with l:w ratio c. 1, appressed, inserted in
middle or more often distal third, slightly convex, ±flat
towards apex, faintly 1 -nerved or with venation obscure,
glabrous or with a few hairs distally, dull brown. Calyx

2.5- 4.5 mm long, glabrous or less often hairy, with tube
equal to or slightly longer than upper lobes; upper lobes

1.5- 2 mm long, 2.5-3.5 mm wide, expanded beyond
lateral angle by 0.3-1 mm; lateral angle acute or
acuminate; sinus 1-1.5 mm deep; lower lobes 0.6-1 mm
long, c. 0.6 mm wide, with lateral lobes flat; standard
to c. 12 mm long, slightly longer than wings and keel;
adaxially yellow with a red flare, with throat generally
not or not fully bisected, abaxially reddish almost
throughout; wings c. as long as keel, c. 2.5 mm wide,
purplish brown, sometimes yellowish near apex, also
variously streaked red proximaily and ventrally; keel
c. 3 mm wide, red throughout; anthers c 0.4 mm long
post-dehiscence; ovary glabrous or rarely with hairs
along lower suture, commonly 4-ovulate; style 3-4 mm
long. Pods: stipe 3-5 mm long; body c. elliptic, 10-16
mm long, 6-9 mm wide, glabrous or rarely sparsely hairy
along lower suture; upper margin c. 0.8 mm wide, with

Muelleria

123

Thompson

ridge to c. 0.5 mm high. Seeds 3-4 mm long, 2-3 mm
wide; aril 1.5-2.5 mm long, 1-1.5 mm high, with base
1 -2 mm long, with lobe curving 90-135° (Fig. 4a, i).

Selected specimens from c 250 examined: SOUTH
AUSTRALIA: W side of Mt Burr golf course, P.Gibbons 25.
4.X.1981 (AD, MEL); Hundred of Hindmarsh, section 455, c.
25 km NW of Mt Gambier, B.BIaylock 23, 5.ix.l965 (AD); Cave
Range, c. 50 km S of Naracoorte, D.Hunt 476, 26.xi.1961 (AD).
VICTORIA: Gippsland Hwy, c. 2 km SE of Cranbourne, T.B.Muir
1264. 29.ix.1960 (MEL); Rotamah Island, The Lakes National
Park, I.Crowford 477, 1 l.ix.l986 (MEL); N margin of Holey Plains
State Park, M.G.Cornck 10035, 24j(i.l986 (MEL); Five Mile Beach,
Wilsons Promontory National Park, RG.A6e//264&CHerscov/fc/?,
4.xii.1986 (MEL, NSW); Forest Camp Track, Gleneig National
Park, RJ.FIetcher 180, 15.ix.1993 (MEL); Yarram Gap, Grampians
National Park, A.CBeauglehole 30918, 8.ix.1969 (AD, MEL);
Jimmys Creek area. 26 km 5 of Halls Gap PO, A.CBeauglehole
66934, 6j(ii.l979 (MEL). TASMANIA: Track to E-shape Lagoon,
Flinders Island, JS.Whinray 9213, undated (AD, CANB, HO, MEL,
NSW); Waterhouse Reserve, near One Tree Hill, AM.Buchanan
752 7,21.xi.l 983 (HO); NEofRisdon Brook reservoir, ACRozefeWs
1427.3ax.1999 (HO).

Flowering period: Flowers in late July to November.

Distribution and habitat: Occurs in far south-eastern
South Australia, southern Victoria and in Tasmania
(Fig. 5a). An old record label giving West Pymble, in New
South Wales {Hellyer, 1964 NSW) is considered to be an
error. Grows in sandy to loamy soils in heathland, scrub,
woodland and forest.

Notes: Bossiaea cinerea is unique in having leaves
in which the leaflet-articulation is indistinct except in
being marked by a spur. Other species that develop a
spur also have a geniculate articulation. Yellow-flowered
forms, i.e., with red markings lacking, have been
occasionally collected, e.g., in Melbourne {S.Rennick 109

MEL) in south-central Victoria, and Edenhope in south¬
western Victoria {Summerhayes, MEL). All populations
of B. cinerea in the Grampians Ranges in south-western
Victoria differ from the typical form in having flowers
with a hairy calyx and with several long hairs on the
lower margin of the ovary. In addition, the leaves of the
Grampians form generally have a more robust and more
elongate apiculum.

Typification: Lee (1970) indicated that a sheet at BM,
with labelling indicating Port Dalrymple was the site of
collection, was presumed to be the holotype. Brown did
not designate a type and is likely to have used material
from both Port Dalrymple and the Derwent River. I do
not consider that Lee effectively lectotypified this sheet
in 1970 by her presumption. I here select this same
sheet, now barcoded BM 000885933, as the lectotype of
B. cinerea. It bears three pieces, with mature fruit evident
on two of them as would be expected for January. CANB
278253, received from BM, has previously been identified
as type material. The label gives Port Dalrymple, which
suggests it may be an isolectotype; however, the single
piece bears flower buds just prior to anthesis. I consider
it impossible for this to have been collected in January
by Brown, so it appears that a labelling error has been
made. Nevertheless the material may still be type
material; it may have been collected by Brown in late
winter while still in southern Tasmania, or it may have
been passed on to him.

Hybridisation: Probable hybrids between B. cinerea
and B. rosmarinifolia have been recorded from the
Grampians Ranges in south-western Victoria {J.Westaway
263 MEL; H.Williomson, xi.l902 NSW; M.Corrick 5317 AO,
MEL).

Figure 5. Distributions of species in Group B. a. Bossiaea cinerea; b. B. rosmarinifolia; c. B. cordifolia; d. 6. kiamensis.

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6. Bossiaea rosmarinifolia Lindl., in T.L.Mitchell,
Three Exped. Australia 2:178 (1838)

Bossiaeo cinerea var. rosmarinifolia (Lindl.) Benth.,
FMusfra/.2:160(1864).

Type: not designated. [Protologue: No locality
information with the description but deduced to be
collected on Mt William in the Grampians Ranges from
other commentary in the text.] Victoria. Mt William,
TlMtchell, vii.1836; probable isotype; K 000278329,
fide A.S.George, in sched., image seen in Kew Herbarium
Catalogue. Type material likely to be located at
CGE also, n.v.

Erect shrubs to c. 3 m high, with inflorescences borne
typically on longer branchlets rather than a regular
series of short side-branchlets; branchlets erecto-
patent, terete, c. 1 mm wide, with hairs 0.5-0.8 mm long;
epicuticular wax often developed. Stipules setaceous,
2-4 mm long, commonly recurved and somewhat
twisted, reddish, soon glabrescent, with venation
obscure; stipule-petiole angle mostly c. 30-60®. Leaves
alternate or a smaller proportion sub-opposite, opposite
or in whorls; petiole 0.5-1.2 mm long; articulation
usually slightly geniculate, with a spur mostly 0.5-1 mm
long; lamina narrow-lanceolate to narrow-linear, 20-30
mm long, 1-5 mm wide, with l:w ratio mostly 8-25, flat
or slightly convex abaxially, markedly discolorous; base
symmetrical, rounded; margin revolute, not undulate,
glabrous; apex narrowly acute; apiculum 1-2 mm
long, mostly pungent, not downcurved; upper surface
smooth, with midrib distinct, but secondary venation
generally obscure, with gland-dotting not evident, soon
glabrescent; lower surface glabrescent. Inflorescences:
axes contracted; bract 0.3-0.5 mm long, c. 0.4 mm wide,
slightly convex abaxially, generally hidden by scales;
pedicel 3-8 mm long, hairy, with hairs often extending
onto receptacle; bracteoles persistent, mostly broad-
ovate, 0.3-0.8 mm long, with l:w ratio c. 1, appressed,
inserted in distal third, slightly convex, with apex flat,
with venation obscure, sparsely hairy distally, dull
brown. Ca/yx 3-4.5 mm long, glabrous, with tube equal
to or slightly longer than upper lobes; upper lobes 1 -2.5
mm long, 2.5-3 mm wide, expanded beyond lateral
angle by 0.3-1 mm; lateral angle acute or more often
acuminate; sinus c. 1 mm deep; lower lobes 0.6-1 mm
long, c. 0.5 mm wide, with lateral lobes flat; standard
to c. 12 mm long, slightly longer than wings and keel,

adaxially yellow with a red flare, abaxially similar or
flushed red over most of surface; wings c. as long as
keel, c. 2.5 mm wide, yellow except for red or brownish-
red marks proximally or in lower half; keel c. 3.5 mm
wide, red tthroughout; anthers 0.5-0.6 mm long post¬
dehiscence; ovary glabrous except for a few long hairs
commonly present in distal half and on lower margin,
3- or 4-ovulate; style 3-4 mm long. Pods: stipe 3-5 mm
long; body c. elliptic, 10-15 mm long, 6-8 mm wide,
glabrous or rarely with a few persistent hairs on lower
margin; upper margin c 0.8 mm wide, hardly ridged.
Seeds 3-4 mm long, 2-2.8 mm wide; aril c. 2 mm long,
c. 1.2 mm high, with base 1.5-2 mm long, with lobe
curving c. 90° (Fig. 4d).

Selected specimens from c. 30 examined: VICTORIA:
Silverband Rd,Grampians, T.&J.Whaite /554, 31.X.1953 (NSW);
Halls Gap-Dunkeld road, 19.2 km S of Halls Creek, R.CWeston
114, 14.X.1984 {CANB, MEL); Mt Rosea, Grampians, M.E.Phillips
496, 4.xi.1971 (CANB, NSW); Bovine Creek crossing on Halls
Gap-Dunkeld road, Grampians National Park, J.H.Ross 3803,
22.ix.1996 (MEL); Halls Gap, CD'Alton, x.1923 (AD).

Flowering period: Flowers in Septemberand October.

Distribution and habitat: Occurs in the Grampians
Ranges of south-western Victoria (Fig. 5b). Categorised
as rare in Australia (Walsh & Stajsic 2007). Grows in dry
sclerophyll open forest.

Notes: Bossiaea rosmarinifolia is immediately
distinguished from other eastern Australian species
by the high length to width ratio of its leaves. It is also
distinguished from B. cinerea, probably its closest
relative, by its longer petioles, geniculate articulation,
smooth leaf-lamina, and standard and wing petals that
are less extensively marked red or purplish-brown. The
flower-bud often has a very pronounced beak, which
is formed from the filiform apices of the calyx-lobes.
It usually has a few hairs nearer the distal end of the
lower suture of the ovary and these hairs can persist
until the developing fruit is more or less a mature size.
These hairs distinguish it from the other members of the
subgroup except for the Grampians form of B. cinerea.

Typification: Type material is likely to be housed
at CGE, where Lindley's herbarium is housed, but
unfortunately this has not been verified at this time,
t have seen an image of K 000278329 which is labelled
as Mt William, July, 37 16 S, 142 Va E, New South Wales,
Mitchell's Expedition 1835. No. 256. It was annotated

Muelleria

125

Thompson

as probable isotype by Alex George in 2005. The date
on the label is a mistake as the year of Mitchell's third
expedition was 1836. The origin of another piece on the
same sheet, K 000278330 is unclear from the label.

Hybridisation: Probable hybrids between 6 . dnerea
and B. rosmarinifolia have been recorded from the
Grampians Ranges in south-western Victoria {J.Westaway
263 MEL; H.Williamson, xi.1902 NSW; M.Corrick 5317 AD,
MEL).

7. Bossiaea cordifolia Sweet, FL Australas,
(Sweet): 20, pi. 20 (1827)

Type: not designated. [Protologue:'... raised from seed,
sent by Mr. Henchman's Collector, Mr. William Baxter,
who collected them on the south coast of New Holland
.1] Holotype: pi. 20 in FL Australas. (Sweet): 20 (1827).
Epitype (here selected): New South Wales. Pambula,
H.Forde, x.1905: NSW 43671.

Bossiaea dnerea sensu G.Bentham, FL Austral. 2: 160
(1864), and subsequent Australian authors, pro parte,
non sensu stricto.

Erect shrubs to c. 3 m high, with inflorescences borne
usually on longer branchlets rather than a regular series
of short side-branchlets; branchlets erecto-patent to
almost spreading, terete, c. 0.5 mm wide, with hairs
0.3-0.8 mm long; epicuticular wax not developed.
Stipules narrow-triangular to filiform, 1-3 mm long,
recurved or deflexed, reddish, hairy at first, with
venation obscure; stipule-pair somewhat adjacent,
forming an angle of c. 30-140° with each other;
stipule-petiole angle not generally measurable due to
deflexing. Leaves variously arranged along a branch,
mostly alternate, but also c. opposite or in whorls;
petiole 0.3-0.8 mm long; articulation obscure; lamina
triangular-ovate, 5-12 mm long, 2-7 mm wide, with
l:w ratio mostly 1.2-2, but occasionally up to 4, convex
laterally, markedly discolorous; base symmetrical,
cordate, truncate or broadly rounded; margin recurved
or slightly revolute, often undulate, occasionally with
a few hairs; apex narrowly acute; apiculum 1-2 mm
long, pungent, not downcurved; upper surface smooth
or minutely tuberculate, with venation generally
slightly raised, glabrous or sparsely hairy; lower surface
commonly glabrous except for veins, sometimes
hairy throughout. Inflorescences: axes contracted or to

c. 2 mm long; hairy; bract c. 0.5 mm long, c. 0.3 mm
wide, slightly convex; pedicel 3-15 mm long, glabrous,
or occasionally sparsely hairy proximally; bracteoles
persistent, variously shaped, 0.3-0.6 mm long, with l:w
ratio 0.5-1, ±appressed, inserted in distal third mostly,
slightly convex, with apex flat or slightly recurved, with
venation obscure, glabrous, dull brown. Calyx 3-4 mm
long, glabrous, with tube longer than upper lobes;
upper lobes 1.5-2.5 mm long, 2.5-3.2 mm wide, often
expanded beyond lateral angle by up to c. 0.5 mm;
lateral angle acuminate; sinus c. 1 mm deep; lower lobes
c. 1 mm long, c. 0.6 mm wide, with lateral lobes ±flat;
standard to c. 11 mm long, slightly longer than wings
and keel, adaxially yellow with a red flare, abaxially often
red over much of surface; wings c. 2.5 mm wide, yellow,
sometimes with a small red mark proximally; keel
3-3.5 mm wide, red ±throughout: anthers 0.5-0.6 mm
long post-dehiscence; ovary glabrous, 3- or 4-ovulate;
style 3-4 mm long. Pods: stipe 3-7 mm long; body
c. elliptic, 15-20 mm long, 5-7 mm wide, glabrous;
upper margin c. 0.8 mm wide, with ridge 0.5 mm high.
Seeds 3-4 mm long, 2-2.5 mm wide; aril 1-2 mm long,
1-1.5 mm high, with base c. 1.2 mm long, with lobe
curving 60-120° (Fig. 4b, f-h).

Selected specimens from c. 50 examined: NEW SOUTH
WALES: c. 2 km W by track from Leonards Island, NE of Eden,
D.FAIbrecht 998, 26.ix.1984 (MEL, CANB); opposite aerodrome,
Merimbula, E.F.Constable 5494, 3>i.1964 (CANB, NSW); 3 km
N of Merimbula, on Merimbula-Tathra road, T.B.Muir 5091,
26.viii.1973 (MEL); junction of Chipmill Rd and road to Boyd's
Tower, M.G.Corrick6030, 18.ix.l 978 (CANB, HO, MEL). VICTORIA:
5 km WNW of Lavers Hill PO,A.C.Beouglehole67375, 19.xii.1979
(MEL); c. 5 km 5 of Chappie Vale, H.IAston 814, 16.xi.l 960 (MEL);
Black Range, EAshby, xi.1937 (AD). TASMANIA: Rocky Cape,
L.Richley, 13.X.1975 (HO); Exploration Creek, Newhaven Track,
AM.Buchanan 15452, 29.vi.1999 (HO); Lake Ashwood, 6 km
NE of Strahan, A.E.Orchard 5739, 6.xii.1981 (AD, HO); 5 km S of
Marrawah, AMBuchanan 14003, 4.X.1995 (CANB, HO).

Flowering period: Flowers from late winter to early
summer.

Distribution and habitat: Occurs in far south-eastern
New South Wales, the Otway Ranges of south-western
Victoria and in western Tasmania (Fig. 5c). Grows in open
forest and heathland.

Notes: Bossiaea cordifolia has a distinctive stipule
orientation. Stipules are inserted somewhat adjacent to
one another, rather than on opposite sides of the leaf

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Vol 30(2) 2012

Eastern Bossiaea

attachment point, and they tend to become deflexed
with age (Fig. 4f). The angle formed by the deflexed
stipules varies from about 30° to 140°, and often they
appear to be connected via a slender rim. In other
species stipules do not usually become deflexed, even
though they may be strongly reflexed, and they are
always inserted opposite each other. Bossiaea cordifolia
can also be distinguished from other two species In
the Cinerea subgroup by the leaflets which have an
obscure articulation and are more triangular-ovate
and with a lower length:width ratio. Its longer, more
pungent leaf-apiculum, glabrous pedicels, and yellow
wing petals usually distinguishes it from B. cinerea. It is
geographically well-separated from both B. cinerea and
B. rosmarinifolia.

A sterile specimen from Nelson Bay River on the
west coast of Tasmania {F.EDavies 1153 CANS) has
leaves intermediate between those of B. cordifolia and
B. cinerea. Further collections from this area are desirable.

Typ/ficaf/on: The holotype illustration is recognisable
as likely to be B. cordifolia rather than B. cinerea based on
the leaflet shape and the yellow wing petals. However,
to make the application of the name more certain, I here
select an epitype, H.Forde, x.1905, NSW 43671, for the
holotype illustration.

The Kiamensis subgroup

8. Bossiaea kiamensis Benth., FL Austral. 2:158
(1864)

Type; [Protologue:'N, S. Wales. Near Kiama, lllawarra,
Backhouse'.] New South Wales. Near Kiama, lllawarra,
J.Backhouse, date unknown; lectotype (here selected):
K 000278246, image seen in Kew Herbarium Catalogue;
isolectotype: K 000278247, image seen in Kew
Herbarium Catalogue.

Erect shrubs to c. 3 m high, with inflorescences borne
on longer branchlets or on a regular series of short
side-branchlets; branchlets erecto-patent, mildly
compressed at first, c. 1 ’mm wide, with hairs 0.2-0,5
mm long, glabrescent; epicuticular wax sometimes
developed. Stipules narrow-triangular, 1.5-4 mm long,
erect, red-brown, gradually glabrescent, 1-3-nerved;
stipule-petiole angle 30-80°. Leaves opposite; petiole
1-1.5 mm long; articulation strongly geniculate, with
a spur 0.3-0.8 mm long; lamina narrow-elliptic, 10-35

mm long, 2-7 mm wide, with l:w ratio 2-6, or rarely to
c. 10, flat or gently convex each side of midrib, markedly
discolorous; base symmetrical, rounded; margin almost
flat to recurved, glabrous, minutely knobbly; apex
subacute to acute; apiculum 0.3-1 mm long, generally
brittle, pointing forwards; upper surface smooth, with
venation raised, brochidodromous, glabrous; lower
surface glabrescent. Inflorescences: axes contracted
or more often to c. 2 mm long, densely hairy; bract
0.5-1 mm long, c. 0.5 mm wide, gently convex; pedicel
3-7 mm long, hairy or glabrous, usually wrinkled
longitudinally below receptacle; bracteoles persistent,
ovate, 0.4-1 mm long, with l:w ratio 1-2, ±appressed,
inserted in distal half, convex, nearly flat at apex, with
venation obscure, glabrous or with a few medial hairs,
mid-brown or red-brown. Calyx A-5 mm long, glabrous,
with tube as long as or shorter than upper lobes; upper
lobes 2-3 mm long, 3-4 mm wide, expanded beyond
lateral angle by 1-2 mm; lateral angle acuminate; sinus
1-2 mm deep; lower lobes 1-1.5 mm long, c 0.5 mm
wide, with lateral lobes convex; standard to c, 12 mm
long, slightly longer than wings and keel, adaxially
yellow with a red flare, with throat bisected, abaxially
generally red, with pale radiating nerves medially;
wings as long as or marginally longer than keel, 2-3 mm
wide, light purplish-brown with red streaks, sometimes
grading to dirty yellowish distally; keel 3-4 mm wide, red
throughout; anthers c. 0.5 mm long post-dehiscence;
ovary glabrous, 3-ovulate; style c. 3 mm long. Pods: stipe
3-4 mm long; body c. elliptic, 10-15 mm long, 7-8 mm
wide, glabrous; upper margin c. 0.7 mm wide, with ridge
to c. 0.5 mm high. Seeds 3-3.5 mm long, 2-2.5 mm wide;
aril 1.5-2.5 mm long,c. 1 mm high, with base 1-1.5 mm
long, with lobe curving 90-130° (Fig. 4c, e).

Selected specimens from c. 90 examined: NEW SOUTH
WALES: The Castle. Budawang Range, I.R.Teiford 237,
22.ix.1967 (CANB); car park at end of Tin Mine Rd (off 12 Mile
Rd), Morton National Park, K.LMcDougall 961, 5.ix.2001 (MEL);
Clyde Mountain, O.D.Evans 1703, 15.ix.l926 (CANB); Budderoo
National Park, Barren Grounds Nature Reserve, F.E.Davies 409
& IMulcahy, 7.xii,1987 (CANB, NSW); Round Hill, c. 5 km S of
Sassafras, SW of Nowra, E.F.Constable, 20.ix.l 961 (MEL, NSW).

Flowering period: Flowers in September and October.

Distribution and habitat Occurs from Kiama south
to Batemans Bay in south-eastern New South Wales, on
near-coastal slopes and mountains (Fig. 5d). Grows in
open forest.

Muelleria

127

Thompson

Notes: Bossiaea kiamensis is readily identified by its
regularly opposite leaves, a strongly discolorous leaflet
which is much longer than wide, and a petiole that is
strongly spurred at the articulation (Fig. 4e). Apart from
opposite leaves, B. kiamensis has a number of other
features linking it to the Cordigera subgroup, including
the retrorsely-directed lateral angle of the upper calyx-
lobes and the short axis on which inflorescence scales
and inflorescences are raised. Pedicels in B. kiamensis are
stouter and fleshier than those of the Cinerea subgroup
and have conspicuous decurrencies below the
bracteoles. Other distinctive features of B. kiamensis are
the convex lower calyx-lobes and verrucose branches
and branchlets. The verrucosities become exposed as
the indumentum is lost. Two inflorescences, one per
axil, are frequently developed at a node. In contrast, in
B. cordigera and R lenticuloris (Cordigera subgroup), the
other two species with opposite leaves, an inflorescence
usually only develops in one of the axils.

7yp/ficaf/on: There is one sheet available for viewing
in the Kew Herbarium Catalogue containing type
material of R kiamensis. I believe all the pieces on the
page to be from the original collection by Backhouse
and would have been seen by Bentham.The specimens
bear flowers. The material is split into two groups
(two barcode identifiers), with three pieces associated
with a Herbarium Hookerianum stamp designated as
K 000278246, and two pieces associated with a
Herbarium Benthamianum stamp designated as
K 000278247. The label associated with K 000278246
matches the protologue better and I therefore choose it
as the lectotype of R kiamensis. Lee in (1970) indicated
that A.B. Court had seen 'the Holotype'; however, it is
unclear whether Lee was referring to the sheet described
above or to another sheet that I have not seen.

Group C

Prostrate or low-growing subshrubs or weakly erect
shrubs; branchlets In 1 or 2 regular series, short, widely
divergent to spreading; terete, with decurrencies absent
or poorly developed. Stipules relatively narrow, brown or
reddish, often with apex filiform, commonly recurved.
Leaves small, not or not much longer than broad,
with articulation sometimes obscure, with apiculum
sometimes slender and recurved. Inflorescences: axes
with scales 2; bracteoles persistent, 1-2 times longer

than wide, sometimes divergent, generally inserted
beyond mid-pedicel. Ca/yxwith upper lobes broadening
from the base, broader than long, mostly expanded well
beyond lateral angle; standard often completely reddish
or brownish abaxially; keel with red marking restricted
to distal half. Pods commonly narrow-oblong. Seeds
often 4 or more per pod, small; aril with a short base and
strongly arched lobes (Fig. 6).

Group C contains five species, and occurs in south¬
eastern Queensland, eastern New South Wales,
southern Victoria, and Tasmania (Fig. 7). It comprises
two well-defined subgroups, the Cordigera subgroup
and the Buxifolia subgroup. Generally speaking, it
is similar to Group B in stipule, bracteole and calyx
morphology and in having markedly discolorous leaves.
The Cordigera subgroup has similarities to B. kiamensis
in particular. The Buxifolia subgroup has similarities to
group D and to the Brownii subgroup of Group E.

The Cordigera subgroup (species 9 & 10) differs from
all other eastern species in its branching pattern and in a
combination ofleafcharactersdeavesareopposite, small,
as broad as long, and with slender petioles (0.1-0.2 mm
in diameter; Fig. 6e, f). Elongation of the inflorescence
axis below the inflorescence scales is typical in this
subgroup. Compared to the Buxifolia subgroup they
have pods with longer stipes, larger anthers, and upper
calyx-lobes with the lateral angle pointing retrorsely.The
shapes and relative sizes of calyx-lobes in this subgroup
as well as that of R decumbens in the Buxifolia subgroup
are reminiscent of the morphology seen in Platylobium.

The Buxifolia subgroup (species 11-13) has bracteoles
that are distinctive in tending to be inserted somewhat
adjacent to each other on the upper side of the pedicel
ratherthan on opposite sides of the pedicel, and in being
more divergent from the pedicel (Fig. 6d). The apiculum
of leaves is often slender, dark and brittle, and often
recurved to slightly hooked (Fig. 6c). Flowers are often
relatively few and sporadic later flowering appears to
occur more often than in other leafy species of eastern
Bossiaea. The pod-stipe is much shorter than the calyx.
The Buxifolia subgroup resembles the Prostrata and
Scortechinii subgroups of Group D in being prostrate
to generally low-growing plants with short-stipitate,
narrow-oblong pods with hairy margins, but also
resembles the Brownii subgroup of Group E in having
almost terete branchlets and leaflets with asymmetric
bases.

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Vol 30(2) 2012

Eastern Bossiaea

Figure 6. Group C. a. Bossiaea decumbens {N.G.Wolsh 1848 MEL); b. 8. lenticularis, {R.Coveny 11912a CANS); c. 8. neoanglica, leaves
{P.CJobson 5203 MEL); d. 8. buxifolia, bracteoles {G.W.Carr 10143 MEL); e. 8. cordigero, leaves and flov/er buds {ASimson 1819 HO);
f. 8. lenticularis, inflorescence shortly after anthesis showing inflorescence axis with elongation below the scales, and reflexed
upper calyx lobes. Arrow is pointing to the two scales and the bract (R.Coveny CANB). Scale barsra, b= 10 mm, c-f=2 mm

The Cordigera subgroup

9. Bossiaea lenticularis Sieber ex DC., Prodr, 2:
117(1825)

Type: [Protologue: 'Sieb! pi. exsic. nov.-holl. n. 425'.]
New South Wales. Location unknown, F.5ieber425, date
unknown; holotype; G-DC, images seen MEL; isotypes:
MEL 668121, MEL 668122, NSW 606082.

Sprawling to erect shrubs to c. 1.5 m high, with
inflorescences borne on a regular series of very short,
side-branchlets which in turn are produced along a
regular series of spreading side-branches; branchlets
spreading, terete, c. 0.4 mm wide, glabrous or sparsely
hairy, glabrescent; hairs 0.1-0.2 mm long; epicuticular

wax absent. Stipules narrow-triangular, 0.4-1 (-3) mm
long, erect, brown, glabrous, with venation obscure;
stipule-petiole angle 30-90°. Leaves opposite; petiole
0.4-1.2 mm long; articulation usually slightly to strongly
geniculate, ridged; lamina circular, oblate, broad-ovate,
occasionally broad-obovate or rhomboid-elliptic, 2-5
(-8) mm long, 2-6{-8) mm wide, with l:w ratio mostly
0.8-1.1, flat, markedly discolorous; base symmetrical,
broadly rounded, truncate or shallowly cordate; margin
slightly recurved, sometimes minutely tuberculate;
apex broadly rounded to subtruncate; apiculum not
developed; upper surface smooth, with venation mostly
obscure, glabrous; lower surface glabrous. Inflorescences:
axes usually 1-5 mm long, with scattered hairs or
glabrous; bract 0.5-1 mm long, c. 0.3 mm wide, convex;

Muelteria

129

Thompson

Key to Group C

1 Leaves opposite_

1: Leaves alternate_

2 Leaflets c. orbicular, with base not or only slightly cordate; branchlets and pedicels glabrous; pedicel

< 5 mm long (New South Wales).9. B. lenticularis

2: Leaflets mostly broad-ovate, with base cordate; branchlets and pedicels hairy, pedicel > 5 mm long

(Victoria and Tasmania)........10. S. cordigera

3 Pedicels < 6 mm long; ovary and pod hairy on valves and marglns,or if sometimes hairs absent or very

few on valves then calyx glabrous; leaflet apiculum typically > 0.5 mm long.13. B. neoanglica

3: Pedicels mostly > 6 mm long; ovary and pod with hairs on margins only; calyx hairy; leaflet apiculum

typically < 0.5 mm long..

4 Upper lobes of calyx 1.5-3 mm longer than lower lobes; keel > 6 mm long; style 2.5-6 mm long.11. B. decumbens

4: Upper lobes of calyx c.0.5 mm longer than lower lobes; keel < 6 mm long; style 1-2 mm long.12. B. buxifolia

pedicel 3-7 mm long, glabrous; bracteoles persistent,
ovate, 0.5-1 mm long, with l:w ratio c. 1-1.5, appressed,
inserted in middle third, mostly beyond mid-pedicel,
convex, with venation usually obscure, glabrous, brown
or red-brown. Co/yx4-5 mm long, glabrous, with tube c.
as long as upper lobes; upper lobes 2-2.5 mm long, 3-4
mm wide, expanded beyond lateral angle by 1.5-2.5 mm;
lateral angle acute or occasionally acuminate; sinus c. 2
mm deep; lower lobes c. 1 mm long, c 0.5 mm wide, with
lateral lobes often slightly convex; standard to c. 12 mm
long, similar in length to wings and keel, adaxially yellow
with a red flare, abaxially often flushed red medially;
wings c. as long as keel, c. 2 mm wide, yellow; keel c 3
mm wide, pale proximally, red distally; anthers c. 0.6 mm
long post-dehiscence; ovary glabrous, 4-6-ovulate; style
2.5-4 mm long. Pods: stipe 5-12 mm long; body elliptic,
rhomboidal or oblong, 10-20 mm long, 5.5-8 mm wide,
glabrous; upper margin c. 0.7 mm wide, with ridge to c.
0.3 mm high. Seeds 2.5-3.5 mm long, 1.5-2 mm wide; aril
1.2-2 mm long, c. 1 mm high, with base c. 1 mm long,
with lobe curving 90-130° (Fig. 6f).

Selected specimens from c. 60 examined: NEW SOUTH
WALES; Mt Wilson, CBurgess, 9.xi.l962 (CANB); Thirlmere,
CBurgess, 1 l.x.1961 (CANB); road to Oakdale State Coal Mine,
c. 5 km NNW of Oakdale, R.Coveny 119i2&P.Weston, 27,ix.l984
(CANB, NSW); South Maroota, c 0.9 km along Paulis Rd from
the Windsor-Vyisemans Ferry Rd, A.E.0rme 176 & R.G.Coveny,
27.X.2001 (BRI, CANB, MEL); Morts Gully, Lithgow, J.LBoorman,
30.X.1914 (NSW); Burragorang Valley, RHCambage 2311,
8.X.1909 (NSW); Grassy Hill, Colo-Putty Rd, E.F.Constable,

7.ix.1948 (NSW); Laughtondale Gully Rd, c. 1 km E of junction
with the Great Northern Rd, Maroota, R.G.Coveny 15522,
22.viii.1991 (AD, BRI, CANB, HO, MEL, NSW, PERTH).

Flowering period: Flowers in spring.

Distribution and habitat: Occurs in near coastal parts
of central eastern New South Wales, including the Blue
Mountains from Howes Valley in the north to Thirlmere in
the south (Fig. 7a).The label on a 1924 collection {Welch.
NSW565910), which gives Tumut as the location, must
be considered doubtful. Grows in sand on sandstone,
often in swampy sites.

Notes: Bossiaea lenticularis is most closely related
to B. cordigera and, like the latter, is readily identifiable
by its divaricate branching, small, opposite leaves
with a circular lamina, relatively slender branchlets
and petioles, and folded upper calyx-lobes. The two
species also have some similarities to species in Group
B, particularly in bracteole, calyx and pod morphology.

10. Bossiaea cordigera Benth. ex Hook.f., Ft.
Tasman. 1 (2): 95, pi. 16 (1856)

Type: [Protologue: 'Widely distributed over the
northern parts of the Island, from the sea-level to
4000 feet, Lawrence, Gunn'. The island is Tasmania,
but was given as V.D.L. (Van Diemen's Land) by the
collectors.] Tasmania. Locality unknown, R.Gunn [171],
date unknown; lectotype (here selected): K 000278235,
image seen in Kew Herbarium Catalogue,

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Residual syntypes (all in Tasmania); Locality unknown,
R.Gunn [171]: K 000278226; Patricks River, R.Gunn [171],
14.xii.1844: K 000278230; York Town, R.Gunn [171],
25.i.1844: K 000278231; George Town, R.Gunn [171],
1842: K 000278232; Lake Arthur, Western Mts, Lawrence:
K 000278236; Circular Head, R.Gunn [171], 1842:
K 000278237; Locality unknown, R.Gunn s.n.:
MEL 651106 (possibly).

Bossiaea hendersonii Regel, Gartenflora 15: 322, pi.
523, 3d, e (1866), as Henderson!. Type: not designated.
[Protologue: Translated from German as 'Cultivated
from the garden of James Booth and Sons, Hamburg,
Germany'.] Holotype: pi. 523,3d, e in Gartenflora 15:322
(1866).

Erect or sprawling shrubs to c. 1.5 m high, with
inflorescences borne on a regular series of very short
side-branchlets which in turn are produced along a
regular series of spreading side-branches; branchlets
spreading, terete, 0.3-0.5 mm wide, sparsely to
moderately hairy; hairs c. 0.2 mm long, wavy to curly:
epicuticular wax not developed. Stipules narrow-
triangular, 0.5-2 mm long, erect, brown, glabrous,
1-nerved or venation obscure; stipule-petiole angle
30-70° Leaves opposite; petiole 1-2 mm long, very
slender; articulation slightly to moderately geniculate,
ridged; lamina ovate to broad-ovate, occasionally
c. circular, 2.5-6 mm long, 2.5-6 mm wide, with l:w
ratio mostly 0.9-1.0, flat or slightly convex laterally,
markedly discolorous; base symmetrical, cordate or
less often broad-cuneate to truncate; margin slightly
recurved, sometimes with hairs persisting, ±smooth;
apex subacute to rounded; apiculum to c. 0.2 mm long,
downcurved, or not developed; upper surface smooth,
with venation obscure, glabrous; lower surface glabrous
or with hairs on midrib. Inflorescences: axes contracted
or to c. 3 mm long, hairy, with a small leaf and stipules
often developed instead of scales, occasionally with 2 or
more nodes below the flower; bract 0.5-1 mm long, c.
0.5 mm wide, slightly convex; pedicel 15-30 mm long,
hairy; bracteoles persistent, ovate, 0.5-1 mm long, with
l:w ratio 1-1.5, ±appressed, inserted at or more often
beyond mid-pedicel, convex, apex nearly flat, 1-nerved,
glabrous, light brown. Calyx 5-6 mm long, glabrous,
with tube shorter than upper lobes; upper lobes 2.5-3.5

mm long, 3-3.5 mm wide, expanded beyond lateral
angle by 2-3 mm; lateral angle acuminate; sinus 2 mm
deep; lower lobes 1-1.7 mm long, c 0.8 mm wide, with
lateral lobes flat; standard to c. 12 mm long, similar in
length to wings and keel, adaxially yellow with red flare,
abaxially brownish-red except towards margins; wings

3- 3.5 mm wide, mostly brownish-red, sometimes yellow
distally; keel c. 4 mm wide, pale proximally, red distally;
anthers c. 0.7 mm long post-dehiscence; ovary glabrous,

4- 8-ovulate; style 3-4 mm long. Pods: stipe 5-8 mm
long: body narrow-oblong, 15-30 mm long, 5-6 mm
wide, glabrous; upper margin c. 0.7 mm wide, hardly
ridged. Seeds 2-2.5 mm long, c. 1.5 mm wide; aril 1-1.2
mm long, c. 0.5 mm high, with base c. 0.7 mm long, with
lobe curving 120-160° (Fig. 6e).

Selected specimens from c. 80 examined: VICTORIA: Farm
Rd, 0.8 km from Morgan Track, Wombat State Forest, J.H.Ross
3693, 13.xii.l995 (CANB, HO, MEL); Buangor Forest Park, 27 km
E of Ararat PO, A.C.Beauglehole 61498, 10.xi.l978 (MEL); Boiler
Swamp Rd, Portland district, €3/ D.Woolcock 1537, 28.xi.1983
(MEL); Lyonville, H.B.WilHamson, i.l916 (MEL); Benwerrin,
Otway Ranges, A.C.F.Gates, xi.1922 (MEL); Domino Rd, c. 6 km
WSW ofTrentham, I.RJhompson 1470, 19.i.2012 (CANB, MEL).
TASMANIA: Tomahawk River, D.LMorris 8171, 12.X.1981 (HO,
MEL); Picketts Plains, A.Moscal 3998, 12.xi.1983 (HO, MEL);
Cradle Mountain Reserve, AM.OIsen, 3.1.1937 (HO); Port Sorell,
W.M.Curtis. x.1944 (AD, HO, MEL); Penstock, A.V.Giblin, xii.1929
(HO).

Flowering period: Flowers in spring and early summer.

Distribution and habitat: Occurs in south-western
and south-central Victoria from Portland east to
Healesville and in northern two-thirds of Tasmania
(Fig. 7b). Categorised as rare in Victoria (Walsh & Stajsic
2007). Grows in open forest, often beside streams or in
damp environments.

Notes: Bossiaea cordigera is mostly closely related
to B. lenticularis q.v. It is readily distinguished from the
other species in Group C by the combination of its
sparse, short and mostly curly hairs, opposite leaves,
ovate, cordate-based leaflets, long pedicels and long
pod-stipes. Calyx-lobes become strongly deflexed
after flowering. A yellow-flowered mutant has been
recorded from Wombat State Forest SE of Daylesford in
south-central Victoria. Several features of 6 . cordigera,
especially the opposite, ovate leaves and the greatly

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131

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enlarged upper calyx-lobes, are reminiscent of species
in the genus Platylobium.

Mature seeds have only been seen from a few
collections. The aril-lobe is relatively slender and
relatively strongly rotated laterally as it arches over. As is
typical of the group the lobe is strongly curved and the
apex often reaches to the seed surface,

Baron Ferdinand von Mueller cited the name
B. horizontalis in First General Report of the Government
Botanist on the Vegetation of the Colony in 1853,
but it appears that the name R horizontalis was never
published. The name appears on several labels of
specimens of R cordigera at MEL suggesting that
Mueller was planning to describe it himself, in which
case Hooker was named as the future author in the
report by mistake, or he was expecting J.D.Hooker to
name the undescribed species as R horizontalis rather
than R cordigera. In 1862, Mueller wrote a description
of R cordigera in Fragmenta Phytographiae Australiae
without making any reference to R horizontalis.

Typification: From a number of similarly suitable
possibilities, K 000278235 is selected here as the
lectotype of R cordigera. It is a large single piece in
the upper right of the sheet, and bears flowers and
immature fruit.

The Buxifolia subgroup

11. Bossiaea decumbens F.Muell., Fragm, 1 (i):
9(1858)

Type: [Protologue: 'Mount Macedon. Dallachi. In
collibus ad amnem Delatite'. Translation: Hills beside
the Delatite River.] Victoria. Mount Macedon, J.DG//achy,
viii.l 849; lectotype (here selected): MEL 18885.

Residual syntypes: Victoria. Mount Macedon, collector
unknown, date unknown: MEL 18886; Victoria. Delatite
River, F.Mueller, 18.iii.l853: MEL 18887, MEL 18888, MEL
18889.

Bossiaea buxifolia sensu G.Bentham, FI. Austral. 2:163
(1864) and subsequent Australian authors, pro parte,
non sensu stricto.

Prostrate to sprawling shrubs to c. 0.3 m high, with
inflorescences typically borne on a regular series of
short side-branchlets; branchlets erecto-patent to
almost spreading, terete, 0.3-0.5 mm wide, mostly
sparsely hairy; hairs 0.2-0.3 mm long; epicuticular wax
not developed. Stipules narrow-triangular, 0.5-2 mm
long, erect or becoming incurved or recurved distally,
red-brown, glabrous, with venation obscure; stipule-
petiole angle 60-90“. Leaves alternate; petiole 0.3 mm
long; articulation obscure; lamina elliptic to broad-
elliptic, 2-5 mm long, 1.5-4 mm wide, with l:w ratio

Figure 7. Distributions of species in Group C. a. Bossiaea lenticularis; b. B. cordigera; c. B. decumbens; d. B. buxifolia; e. R neoangiica.

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Vol 30(2) 2012

Eastern Bossiaea

1.1-1.8, ±flat, markedly discolorous; base often slightly
asymmetrical, slightly cordate to rounded; margin
slightly recurved, glabrescent, smooth; apex rounded
to obtuse, often recurved; apiculum to c. 0.4 mm long,
slender, brittle, recurved; upper surface smooth, with
venation variably raised, mostly soon glabrescent; lower
surface glabrescent. Inflorescences: axes contracted;
bract 0.8-1.5 mm long, c. 0.5 mm wide, flat to slightly
convex; pedicel 10-25 mm long, hairy; bracteoles
persistent, oblong to oblong-elliptic or obovate, 0.7-2
mm long, with l:w ratio 1.5-2, slightly to markedly
divergent, inserted beyond mid-pedicel, mostly in
distal third, fairly flat, sometimes recurved distally, with
margins sometimes recurved, with apex flat or convex,
1-nerved, glabrous, light or red-brown. Calyx 3-5.5 mm
long, hairy, with tube much shorter than upper lobes;
upper lobes 2.5-3.5 mm long, 2.5-3 mm wide, expanded
beyond lateral angle by 1-2 mm; lateral angle subacute
to rounded; sinus 1-2.5 mm deep; lower lobes 1-1.5
mm long, 0.8 mm wide, with lateral lobes flat; standard
to c 10 mm long, slightly longer than wings and keel,
adaxially yellow with a red flare, with throat not bisected,
abaxially brownish-red throughout; wings c. as long as
keel, 2-2.5 mm wide, light purplish-brown throughout;
keel 3-4 mm wide, pale except for pink markings in
distal quarter; anthers c. 0.5 mm long post-dehiscence;
ovary with hairs on margins, sometimes hairs rather few,
6-10-ovulate; style 3-6 mm long. Pods: stipe 1-3 mm
long; body narrow-oblong, 20-30 mm long, 5.5-7 mm
wide, with appressed or spreading hairs on margins,
occasionally glabrescent; upper margin c. 0.6 mm wide,
hardly ridged. Seeds 2-3 mm long, 1.2-1.8 mm wide; aril
1-1.5 mm long, 0.6-0.8 mm high, with base 0.5-0.8 mm
long, with lobe curving 135-180° (Fig. 6a).

Selected specimens from c. 50 examined: VICTORIA:
Wonnangatta Valley, c. 1 km NW of the junction of the
Wonnangatta River and Zeka Creek, D.EAlbrecht 3885,
30.xi.1989 (MEL); Cheshunt-Dandongadale Rd, 4.8 km from
Rose Valley, TJ.Entwisle 1725 & S.Bodsworth, 9.X.1990 (CANB,
MEL, PERTH); S bank of Howqua River, c. 400 m downstream
from Sheepyard Flat, N.G.Walsh 1848, 28.V.1987 (CANB, MEL,
NSW); Eagle Point, Mt Buffalo, J.H.Willis, 20.ii.l963 (MEL);
Tipperary Track, S of Bryces Flat, E side of Sailors Creek, SW of
Hepburn, J.H.Ross 3807, 12.X.1996 (MEL); 3 km SE of Beaufort,
A.C.Beauglehole 61687, 19.xi.l978 (MEL).

Flowering period: Flowers from spring to early
summer.

Distribution and habitat: Occurs in southern Victoria
from Ararat east to Bright (Fig. 7c). Grows in open forest.

Notes: Bossiaea decumbens differs from B. buxifolia in
floral morphology with flowers having a longer calyx
and corolla, upper calyx-lobes that are more expanded
beyond the lateral margin, larger anthers, and a longer
style. The red markings of the keel are also generally
paler, stipules are generally less recurved, and leaflets
are flatter and with margins less recurved. The two
species are geographically separated.

Typihcation: From the type material at MEL, I have
selected MEL 18885 as the lectotype as this sheet is the
only one that contains good examples of the flowers.
The original label specifies the collector as Dallachy;
however the date of collection given in the type details
above is based on a recent annotation. Although also
collected from Mt Macedon, it is uncertain whether
MEL 18886 is an isolectotype. The single piece has a
different look to the pieces of the lectotype. However,
this sheet has also been recently annotated indicating
that the collector was Dallachy and the collection was
in August 1849.

12. Bossiaea buxifolia A.Cunn., in B.Field,

Geogr. Memoirs New South Wales: 348 (1825)

Type: [Protologue: 'Upon rocky, brushy hills'.] New
South Wales. Blue Mountains, ACunn/ng/iam; lectotype:
K 000278436, fide Lee (1970).

Prostrate to weakly erect shrubs to c. 0.5 m high, with
inflorescences typically borne on a regular series of
short side-branchlets; branchlets erecto-patent to
almost spreading, terete, 0.3-0.5 mm wide, sparsely to
moderately hairy; hairs 0.2-0.5 mm long; epicuticular
wax sometimes developed. Stipules narrow-triangular
to setaceous, 1-2 mm long, erect or more often widely
divergent and/or becoming recurved, red-brown, hairy,
glabrescent, with venation obscure; stipule-petiole
angle 60-90°. Leaves alternate; petiole 0.2-0.5 mm long;
articulation obscure; lamina elliptic or c. circular, mostly
2.5-5 mm long, 2-5 mm wide, with l:w ratio 1.1-1.6, ±flat
or sometimes slightly concave proximally, markedly
discolorous; base usually slightly asymmetrical, cordate
or truncate; margin slightly recurved to slightly revolute,
hairy at first, with persistent tubercles; apex rounded to
obtuse, straight or slightly recurved; apiculum hardly

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133

Thompson

developed or to 0.5(-0.8) mm long, setaceous, generally
brittle, mostly recurved, sometimes slightly hooked;
upper surface smooth or minutely tuberculate, with
venation variably raised, glabrescent; lower surface hairy
throughout or glabrous except for veins. Inflorescences:
axes contracted or to c. 3 mm long; bract 0.5-1 mm long,
c. 0.5 mm wide, strongly convex; pedicel (3-)6-20 mm
long, hairy; bracteoles persistent, elliptic, broad-elliptic,
oblong-elliptic, or obovate, 0.6-1.2 mm long, with l:w
ratio 1.2-2, loosely appressed or divergent, mostly
inserted in distal half, flat or slightly convex, sometimes
with margins recurved with apex ±flat, 1-nerved or
with venation obscure, glabrous or hairy, brown or red-
brown. Calyx 3-4.5 mm long, hairy, with tube shorter
than or equal to upper lobes; upper lobes 1.5-2.5 mm
long, 2-3 mm wide, expanded beyond lateral angle by

O. 5-1.5 mm; lateral angle subacute, sometimes minutely
acuminate; sinus 1-1.5 mm deep; lower lobes 1-2 mm
long, 0.6 mm wide, with lateral lobes flat; standard to
c. 9 mm long, c 2 mm longer than wings and keel,
adaxially yellow with a red flare, with throat not bisected,
abaxially dark red throughout; wings c. as long as keel,
c. 2 mm wide, red and purplish, sometimes yellowish
distally; keel c. 3 mm wide, pale proximally, dark red
in distal half; anthers c. 0.4 mm long post-dehiscence;
ovary with hairy margins, 5-12-ovulate; style 1.5-2 mm
long. Pods: stipe 1-2 mm long; body narrow-oblong
or narrow oblong-elliptic, 15-30 mm long, 4.5-7 mm
wide, with appressed or occasionally spreading hairs
on margins; upper margin c. 0.6 mm wide, not ridged.
Seeds 2-3 mm long, 1.2-1.5 mm wide; aril 1-1.2 mm
long, 0.6-0.8 mm high, with base c. 0.7 mm long, with
lobe curving c. 180° (Figs 6d, 1 Og).

Selected specimens from c, ISO examined: QUEENSLAND:
Catchment of Precipice Creek, Precipice National Park,

P. I.Forster 19736, 25.ix.1996 (BRI, MEL, NSW); Kroombit Creek,
SW of Annies Gorge, Kroombit National Park, J.Brushe 665 &
R.Hendry. 31.xii.l996 (BRI); State Forest 665, 4 km SE of Crows
Nest, A.R.Bean 7953 & JJhompson, 13.X.1994 (BRI); Barakula
State Forest. V.Hando 13, 3.X.1978 (BRI). NEW SOUTH WALES:
Tia Falls, 100 m E of picnic area, Oxley Wild Rivers National
Park, LM.Copeland 4478, 2x12010 (BRI, CANB, MEL, NSW);_
Tinderry Mountains, Tinderry Nature Reserve, GStewarT 293
& RWhigham. 13.xi.1984 (CANB, MEL, NSW); c. 14 km from
Delegate toward Bombala, EJ.CarroH, 16.xii.1965 (CANB, MEL);
Intersection of Oellen Ford and Yarralaw Rds, c. 10 km N of
Windellama, I.R.Thompson 1279, 30.ix.2010 (BRI, CANB, MEL).

VICTORIA: Mail-box Gully, near Wulgulmerang Creek, on
road to Deddick, J.H.WilHs, 29jci.l962 (MEL); Yambullah peak
track, 4.8 km E of Mt Coopracambra, N.G.Walsh 1218, x.1983
(MEL); Buchan River at Diggers Hole Track crossing, 6 km SW
of Mt Seldom Seen, SJ.Forbes 3200, 3.xi.1986 (CANB, MEL,
NSW); Providence ponds FFR, A.C.Beauglehole 78751, 22.X.1987
(CANB, HO, MEL).

Flowering period: Flowers from spring to early
summer.

Distribution and habitat: Occurs in south-eastern
Queensland south from Kroombit Tops, in eastern New
South Wales, and in eastern Victoria (Fig. 7d). Grows in
dry sclerophyll forest and woodland.

Notes: Bossiaea buxifoHa is a widespread species
which exhibits a moderate amount of variation in
habit, flower size, pedicel length and pod size and
ovule number. It is not always easily distinguished from
B. neoongfica q.v. and B. decumbens q.v.

Typihcation: There appears to have been several
specimens that Cunningham would have had access
to when describing the species. As Cunningham
did not cite a specimen or location when naming
B. buxifolia, the selection of the three pieces barcoded
as K000278436 by Alma Lee in concert with A.B.Court
(Lee 1970; sheet annotated by Court) as the holotype, in
effect lectotypified this sheet.

13. Bossiaea neoanglica F.Muell., Fragm, 5(32):
106 (1865), as Neo-Anglica

Type: [Protologue:'ln collibus lapidosis Novae Angliae
baud procul ab origine fluvii McLeay's River. C. Moore'.
Translation: In stony hills of New England near the
origins of the Macleay River.] New South Wales. Head
of the Macleay River, C.Moore 131, date unknown;
holotype: MEL 18890; possible isotype; NSW 43644 n.v.,
fide Lee (1970).

Prostrate to weakly erect shrubs to c. 0.5 m high, or higher
when supported, with inflorescences typically borne on a
regular series of short side-branchlets;branchletserecto-
patent to almost spreading, c. terete, 0.3-0.5 mm wide,
moderately hairy; hairs sometimes curly, 0.5-0.8 mm
long; epicuticular wax not developed. Stipules narrow-
triangular to filiform, 2-3.5 mm long, mostly recurved or
almost decurved, red-brown, hairy, glabrescent, faintly
1 -nerved; stipule-petiole angle 60-90^^. Leaves alternate;
petiole 0.5-0.8 mm long, articulation sometimes slightly

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Vol 30(2) 2012

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geniculate and/or slightly ridged, sometimes obscure;
lamina ovate or broad-ovate, 2-8 mm long, 2-7 mm
wide, with l:w ratio mostly 1.2-1.8, flat or sometimes
slightly concave proximally, markedly discolorous;
base mostly asymmetrical, cordate or truncate; margin
slightly recurved to slightly revolute, glabrescent, finally
minutely tuberculate; apex rounded to acute, flat or
recurved; apiculum mostly 0.5-1 mm long, setaceous,
brittle, mostly recurved, often slightly hooked; upper
surface smooth or minutely tuberculate, with venation
variably raised, glabrescent; lower surface generally
hairy. Inflorescences: axes contracted or rarely to
c, 2 mm long; bract 0.7-1 mm long, c. 0.6 mm wide,
convex; pedicel 1-6 mm long, glabrous or hairy;
bracteoles persistent, oblong to elliptic or obovate,

1- 1.2 mm long, with l:w ratio 1.5-2, divergent, inserted
c. mid-pedicel or occasionally more distally, nearly
flat or gently convex, with apex sometimes slightly
incurved, faintly 1-nerved, glabrous, red-brown. Calyx
3-4 mm long, glabrous or hairy, with tube shorter than
or equal to upper lobes; upper lobes 2-2.5 mm long,

2- 3 mm wide, expanded beyond lateral angle by
c. 1 mm; lateral angle subacute or obtuse, sometimes
minutely acuminate; sinus 1-2 mm deep; lower lobes

1- 1.5 mm long, c. 0.7 mm wide, with lateral lobes flat;
standard to c. 8 mm long, similar in length to wings
and keel, adaxially yellow with a red flare, abaxially red
tthroughout; wings c. as long as keel, c. 2 mm wide,
red streaked, variously pale purplish-brown or yellow
distally; keel 2.5-3 mm wide, pale proximally, red
distally; anthers c. 0.4 mm long post-dehiscence; ovary
hairy throughout or on margins, 6-8-ovulate; style

2- 3 mm long. Pods: stipe 1-2 mm long; body oblong,
15-25 mm long, 6-8 mm wide, with long, spreading hairs
on valves and margins or valves sometimes glabrous
(a few hairs usually present early in development): upper
margin c. 0.7 mm wide, not or hardly ridged. Seeds 2.5-3
mm long, 1.5-2 mm wide; aril 1-1.3 mm long, 0.8 mm
high, with base c. 0.6 mm long, with lobe curving c. 180°
(Fig. 6c).

Selected specimens from c. 60 examined: QUEENSLAND:
Kroombit Tops State Forest, 2.1 km S of Locked Gate sign on
loop road to Annies Gorge. IBrushe 680 SiR.Hendry, 30.xii.1996
(BRI); Mt Bangalora, Main Range National Park, PlForster 12229
& G.Leiper, 29.X.1992 (BRI, MEL, NSW); New England Hwy, 8.8.
km S of Crows Nest, A.R.Bean 17310, 26.i.2001 (BRI). NEW
SOUTH WALES; Gibraltar Range, c. 59.5 km NE of Glen Innes,
IBWilliams, xi.1959 (NE); track to Dandahra Falls, Gibraltar

Range National Park, R.G.Coveny 16686 & AJ.Whalen, 19.X.1993
(BRI, MEL, NE, NSW); 60 Foot Falls track, freeway underpass,
[Mittagong], GJ.Chandler 963, 27.ix.1999 (CANB); 0.7 km S of
the S lake picnic area, Thirlmere Lakes National Park via Buxton,
Ay.Slee2302, 18.X.1988 (CANB); Gloucester Tops, c. 59 km from
Gloucester, J.Pulley 711. 11.ii.l971 (CANB); Doughboy Range,
Ebor area, RiVJessup 237,18.xi.1953 (CANB); 1.1 km along track
to Basket Swamp rest area, 16.9 km NE ofTenterfield, Boonoo
State Forest, P.CJobson 5203 & SAMills, 25.X.1997 (MEL, NSW);

11.4 km NNW of Oakdale, P.CJohson 3758, 17.ix.l995 (BRI, MEL,
NSW); c. 1.5 km E of Cobcrofts Rd along Mesa ManagementTrail,
Werrikimbe National Park, c. 60 km SE of Walcha, LM.Copeland
4476, 2.xi.2010 (CANB, MEL, NSW).

Flowering period: Flowers sporadically, but mostly in
spring.

Distribution and habitat: Occurs in south-eastern
Queensland south from Kroombit Tops, and in north¬
eastern and central-eastern New South Wales as far
south as Fitzroy Falls (Fig. 7e). Grows in open forest and
woodland.

Notes: Bossiaea neoanglica is superficially very siniilar
to B. buxifolia but, apart from differences indicated in
key, has longer stipules, leaves that are more markedly
asymmetrical, more strongly discolorous, and with a
usually more distinct articulation (under magnification),
and a usually longer apiculum. The long apiculum is
brittle and on herbarium sheets a high proportion are
reduced in length due to breakage. A form from central-
eastern New South Wales differs fairly consistently from
the type form from north-eastern New South Wales
and south-eastern Queensland in having glabrous or
near-glabrous pedicels and calyces, and ovaries/pods
glabrous or only sparsely hairy on the faces.

Group D

Subshrubs or shrubs, often prostrate; branchlets mostly
compressed, with decurrencies sometimes well-
developed but not winged, sometimes spinescent.
Stipules narrow-triangular, generally erect, sometimes
green, generally ±glabrous. Leaves with articulation
generally markedly geniculate; lamina with gland-
dotting generally evident; apiculum generally
inconspicuous. Inflorescences: axes with scales 2, or
sometimes leafy; inflorescences sometimes somewhat
elongated; bract and bracteoles generally strongly
convex, often striate, bracteoles sometimes caducous,
often relatively slender, inserted in proximal or middle

Muelleria

135

Thompson

thirds. Calyx mostly hairy, with lobes often filiform
apically and with lower lobes as long as or longer than
upper lobes and tube; upper lobes mostly toblong, as
long as or longer than broad, often abruptly broadening
at apex, not expanded beyond lateral angle. Corolla
sometimes almost entirely yellow. Pods short-stipitate,
mostly hairy on margins, sometimes hairy on valve
faces also. Aril mostly small, with base short and lobe
moderately to strongly arched (Fig. 8).

Group D contains seven species divided into three
subgroups. It occurs in south-eastern Queensland,
eastern New South Wales, south-eastern and southern
Victoria, Tasmania, and south-eastern South Australia
(Fig. 9).

The Prostrata subgroup (species 14 & 15) is
distinguished from the Scortechinii subgroup by the
more or less flat leaflet-margins, narrower, more scarious
stipules, more consistently suppressed inflorescence
axes, and the typically glabrous pod-valves.

The Scortechinii subgroup (species 16-18) has
leaflets with recurved to revolute margins, stipules
that are somewhat persistently green at least in part,
inflorescences that sometimes become markedly
elongated, filiform calyx-lobe apices, petals with
relatively little or no red coloration, and hairy pod-
valves, The stipules in this subgroup are similar in form
to the much larger stipules of R stephensonii (Group E).

Figure 8. Group D. a. Bossiaea nummularia {A.A.Hami!ton NSW 43654); b. B. obovata (L.M.Copeland 4483 MEL); c. B. scortechinii
[LMCopeland 4493 MEL); d. B. prostrata {I.R.Thompson s.n., 6.xi.2010 MEL); e. B. prostrata, scale, bract and bracteoles [RMSmith
59/247 MEL); f. B. scortechinii, pod {LMCopeland4493 MEL); g. B. obcordata, spinescent branch {N.M.Taws527 CANB);
h. B. tasmonica, sub-spinescent branch; i. B. obcordata {M.E.Phillips 3 CAUB);j. B. tasmanica {Leaman, 8.xii.2010 MEL).

Scale bars:a-d = 10 mm, e,f, i,j = 5 mm, g, h = 2 mm.

136

Vol 30(2) 2012

Eastern Bossioea

The Obcordata subgroup (species 19 & 20) is
distinguished from all other eastern species by its
spinescent or subspinescent branchlets* There are
several Western Australian species of Bossiaea with
spinescent branchlets, and there is generally not a lot
of difference, with the exception perhaps of their woolly
keel-apices and more setaceous stipules, between these
western and eastern species, Bossiaea tasmanica has
more features in common with the other subgroups of
Group D, while B. obcordata has more features that link
it to Group E. The branching pattern in B. obcordata is
also similar to that seen in the Cordigera subgroup of
Group C.

The Prostrata subgroup
14. Bossiaea nummularia Endl., in S.L.Endlicher
& E.FenzI, Nov. Stirp. Dec. 3:22 (1839)

Type: not designated. [Protologue: 'Colitur in horto
Hugeliano'. Translation; Cultivated in gardens of C. von
Hugel, Vienna, Austria.] Probable holotype: W 0031366,
image seen in Naturhistorisches Museum Wien, Virtual
Herbaria.

Bossiaea prostrata pro parte sensu A.T.Lee, Contr.
New South Wales Natl Herb. 4(3): 102 (1970); A.T.Lee &
J.Thompson, FI. New South Woles 101(2): 106 (1984);
T.AJames & GJ.Harden, FI. New South Woles, rev. edn 2:
515(2002).

Prostrate or decumbent subshrubs to c. 0.2 m high, with
inflorescences borne on branchlets of various lengths,
sometimes on a regular series of short side-branchlets;
branchlets erecto-patent, slightly to moderately
compressed, 0.6-1 mm wide, without decurrent ridges,
moderately hairy; hairs c. 0.3 mm long; epicuticular
wax sometimes developed. Stipules narrow-triangular,
sometimes filiform distally, 1-2.5 mm long, erect or
slightly divergent, sometimes partly herbaceous at
first, becoming brown, glabrous or hairy, 1-nerved;
stipule-petiole angle 45-90°. Leaves: petiole 0.5-1 mm
long; articulation slightly to moderately geniculate,
sometimes obscure, not ridged; lamina broad-elliptic,
or less often suborbicular or slightly broad-ovate,
2-12 mm long, 2-6 mm wide, with l:w ratio mostly
1-2, ±f]at, mildly discolorous; base symmetrical,
broadly rounded to slightly cordate; margin flat or

Key to Group D

1 Branchlets not spinescent.2

1: Branchlets spinescent or subspinescent (tapering to a blunt point).6

2 Ovaries and pods with hairs restricted to sutures (rarely with hairs all over in B. prostrata); leaflet margin

nearly flat; standard extensively marked red abaxially; stipules generally brown;.3

2: Ovaries and pods with hairs all over; leaflet margin recurved to revolute; standard yellow or flushed pink

abaxially; stipules commonly substantially green; (northern New South Wales and Queensland).4

3 Petioles £ 1 mm long; bracteoles generally persistent, mostly inserted in middle third of pedicel;

standard-limb ± solidly red abaxially.-. nummularia

3: Longest petioles > 1 mm long; bracteoles generally caducous, inserted in proximal third of pedicel;

standard-limb with long pale streaks interrupting the red markings abaxially. 1 5. B. prostrata

4 Leaflets predominantly obovate; pedicels mostly < 5 mm long.18* obovata

4: Leaflets predominantly narrow-oblong to narrow-elliptic; pedicels mostly > 5 mm long.5

5 Petioles > 1 mm long; leaflets with l:w ratio mostly < 3; bracteoles narrow-elliptic; pods 6-8 mm wide.16. B. dasycarpa

5: Petioles £ 1 mm long; leaflets with l;w ratio mostly > 3; bracteoles very narrow-oblong;

pods 4-5.5 mm wide. .. B.scortechinii

6 Prostrate or low-growing shrubs; calyx hairy; keel-apex greenish-yellow, sometimes tinged pink;

pod valves hairy (Tasmania).19. B. tasmanica

6: Erect shrubs; calyx glabrous or nearly so; keel-apex dark red; pod valves glabrous (mainland states).20. fi. obcordata

Muelleria

137

Thompson

slightly recurved, glabrescent, sometimes sparsely and
minutely tuberculate; apex broadly rounded to obtuse,
or occasionally acute, sometimes minutely acuminate,
sometimes downcurved; apiculum to c 0.3 mm long,
brittle, pointing forward to moderately downward;
upper surface smooth, with venation generally obscure,
with gland-dotting usually evident, with appressed hairs
or glabrescent; lower surface gradually glabrescent.
Inflorescences: axes contracted or rarely to c. 1 mm long,
inflorescences rarely with a short rachis developed when
2-flowered; bract 1-1.5 mm long, 0.6-0.8 mm wide,
moderately convex; pedicel 3-15 mm long, commonly
> 8 mm long, hairy, with hairs commonly appressed;
bracteoles generally persistent, ovate to narrow-ovate
or narrow-oblong, 1-2 mm long, with l:w ratio 1.5-3,
loosely appressed or divergent, inserted in middle third
of pedicel, or occasionally more proximally, moderately
convex. 1-nerved or with venation obscure, usually
hairy, especially distally, red-brown. Calyx 3.5-4.5 mm
long, hairy, sometimes near-glabrous proximally, with
tube c. equal in length to lobes; upper lobes 1.3-2.5 mm
long, 1.5-2 mm wide; lateral angle acute or acuminate;
sinus 1-1.5 mm deep; lower lobes 1.3-2.5 mm long,
c. 0.8 mm wide, with lateral lobes flat; standard to c. 9
mm long, slightly longer than wings and keel, adaxially
yellow with a red flare, abaxially ±entirely red; wings c. as
long as keel, 2-2.5 mm wide, brownish-red throughout
or nearly so; keel 2.5-3 mm wide, pale proximally, red
distally; anthers c. 0.5 mm long post-dehiscence; ovary
with hairy margins, 6-8-ovulate; style 1.5-2.5 mm long.
Pods: stipe 1-3 mm long; body narrow-oblong, 20-30
mm long, 6-7 mm wide, with margins hairy, rarely with
a few hairs on valves; upper margin c. 0.7 mm wide, not
ridged. Seeds 2.5 mm long, 1.5 mm wide; aril c. 0.8 mm
long, c. 0.5 mm high, with basec. 0.4 mm long, with lobe
curving 150-180° (Fig. 8a).

Selected specimens from c. 25 examined: NEW SOUTH
WALES: Parramatta, W,Woolls (MEL); Whalan, R.Coveny 11873
& S.Coodwin. 13.ix,1984 (MEL, NSW); Duck River, Clyde,
A.A.Hamilton, ix.l914 (NSW); Marayong, P.Hind s.n., ix.l967
(NSW); Lumley Rd, 1 km N of Jacqua Rd, c. 30 km direct S of
Goulburn, I.RJhompson 1331, 24.xi.2010 (CANB, MEL, NSW);
near Crookwell,/M.Ooivnes, 25j<i.1983 (NSW); 1.5 km E of Gara
River crossing on Armidale-Grafton road, BJ.Wallace 012/86 &
PGAbell, 22M986 (NSW).

Flowering period: Flowers from August to October.

Distribution and habitat: Occurs in south-eastern
and central-eastern New South Wales, mostly south of
Goulburn and in the Sydney region, with an outlier in
north-eastern New South Wales near Armidale (Fig. 9b).
Grows in woodland and open forest.

Notes: Bossiaea nummularia is superficially similar
to B. prostrata, but differs from that species by having
smaller leaves with a more uniform shape, shorter
petioles, a less geniculate leaflet-articulation, narrower
bracts, and more persistent bracteoles that are inserted
mostly in the middle third rather than proximal third
of the pedicel. In addition, the abaxial surface of the
standard is a solid red rather than with long, radiating
pale streaks as in B. prostrata. Branchlets are generally
slightly less compressed than those of B. prostrata and
can become more extensively covered with epicuticular
wax. Bossiaea nummularia can develop a series of
short side-branchlets similar to that seen in B, buxifolia,
whereas B. prostrata does not generally do so.

The distribution of B. nummularia overlaps that of
B. prostrata and there are several mixed collections
containing these two species. Bossiaea nummularia also
approaches B. buxifolia in some respects and these two
species also overlap in distribution. Bossiaea buxifolia
differs in having branchlets more nearly terete, stipules
darker red, more slender and more recurved, an always
obscure leaflet-articulation, a slightly asymmetric leaflet'
base, more recurved leaflet-margins, and bracteoles that
are inserted more distally, more divergent and tending
to be inserted towards the same side of the pedicel.

Typihcation: The evidence from probable type
material of B. nummularia housed at W, and the
description in the protologue is sufficient I believe to
assign the name B. nummularia to the taxon described
above. The main anomaly is the apparent colour of
the petals in the holotype material which appear to
be all yellow rather than with red markings. Bossiaea
linnaeoides is another potential synonym for this
taxon, and this name was published earlier than
B. nummularia; however, it is unclear from the description
of B. linnaeoides to which taxon Don was referring, and
no specimens or illustrations are known to the author.

15. Bossiaea prostrata R.Br., in W.T.Aiton,

Hortus Kew„ 2nd edn, 4:268 (1812)

Type: not designated. [Protologue: 'Native of New
South Wales. Robert Brown, Esq'.]: New South Wales-

138

Voi 30(2) 2012

Eastern Bossioea

Port Jackson, R.Brown, 1803; lectotype: BM 000885986,
fide Lee (1970).

Residual syntypes: MEL 1528713, MEL 1528712,
NSW171069 [These sheets are either mixtures of
B. prostrota and B. nummularia or are entirely of
B. nummularia.]

Prostrate or decumbent subshrubs to c. 0.2 m high,
with inflorescences borne on branchlets of various
lengths, but not typically on a regular series of short
side-branchlets; branchlets erecto-patent, moderately
compressed, 0.2-0.8 mm wide, sometimes with
decurrent ridges, mostly sparsely hairy, glabrescent;
hairs c. 0.3 mm long; epicuticular wax sometimes weakly
developed. 5f/pu/es narrow-triangular, 0.8-1.5 mm long,
with l:w ratio 1.5-2, erect, herbaceous at first, becoming
brown, glabrous, 1-nerved or obscurely multinerved;
stipule-petiole angle 30-60°. Leaves: petiole 1-5
mm long; articulation strongly geniculate, generally
slightly ridged; lamina circular, oblong to elliptic, or
ovate to lanceolate, 4-15(-25) mm long, 2-15 mm
wide, with l:w ratio mostly 1-3, generally flat or convex
each side of midrib distally, mildly discolorous; base
symmetrical, truncate to slightly cordate; margin ±flat,
glabrescent, ±smooth or tuberculate; apex broadly
rounded to obtuse, or occasionally acute, sometimes
minutely acuminate; apiculum to c. 0.3 mm long, brittle,
pointing forwards or down; upper surface smooth,
with venation commonly slightly raised, with gland-
dotting usually evident, glabrous or glabrescent; lower
surface glabrescent. Inflorescences: axes contracted
or to c. 2 mm long; bract 1.5-2.5 mm long, 1-1.5 mm
wide, strongly convex abaxially, generally conspicuously
striate, often caducous; pedicel (2-)5-20 mm long, hairy,
with hairs commonly spreading; bracteoles caducous
before or after anthesis, narrow-elliptic, narrow- to very-
narrow oblong, oroblanceolate, 1.5-3.5 mm long, with
l:w ratio 1.5-5, loosely appressed or divergent, inserted
mostly in proximal third, often close to base, strongly
convex, 4-8-nerved, usually hairy, especially medially
and distally, red-brown. Calyx 3-4.5 mm long, hairy,with
tube equal to or shorter than lobes; upper lobes 2-3
mm long, 1.5-2 mm wide, abruptly broadening at apex;
lateral angle acuminate; sinus 0.5-1 mm deep; lower
lobes 1.5-3 mm long, filiform distally, c. 0.8 mm wide,
with lateral lobes flat; standard to c. 9 mm long, slightly
longer than wings and keel, adaxially yellow with a

red flare, abaxially red except for pale radiating lines;
wings c. as long as keel, 2-2.5 mm wide, brownish-red
throughout or giving way to yellow distally; keel 2.5-3
mm wide, pale proximally abruptly giving way to red
in distal half; anthers c. 0.4 mm long post-dehiscence;
ovary with hairy margins, 8-10-ovulate; style 2-3 mm
long. Pods: stipe 1-3 mm long; body narrow-oblong,
20-30 mm long, 5-7 mm wide, mostly with one or both
margins hairy, rarely glabrous, rarely hairy on valves;
upper margin 0.7 mm wide, flat to gently convex,
occasionally with a small sutural ridge. Seeds 2-3 mm
long, 1-1.8 mm wide; aril 0.8-1.2 mm long, 0.5-0.8
mm high, with base c. 0.5 mm long, with lobe curving
150-180° (Fig. 8d-e).

Selected specimens from c. 300 examined: SOUTH
AUSTRALIA:6.8kmWSWofTerka,Mt Remarkable National Park,
Flinders Ranges, W.R.Telfer&COatsBSW4-4747, 22.xi.1999 (AD);
Mt Magnificent Conservation Park [Southern Lofty], B.M.Grivell
140, 14.X.1984 (AD); Mt Beevor, [Murray], D.E.Murfet 4225 &
R.L.Toplin, 8.xi.2002 (AD); Hindmarsh Waterfall Rd, road junction
c. 15 km N ofVictor Harbor, J.Z.Weber617, 24.X.1967 (AD); Big
Heath National Park, c. 1 km NW of cairn, C.R.AIcock 2887,
3.xi.l969 (AD). QUEENSLAND: Widgee Mountain, P.I.Forster
12121 & P.Machin, 25x1992 (BRI); Kroombit Tops, State Forest
316,48 km E of Biloela, N.absonTOI433, 17.ix.l988 (BRI). NEW
SOUTH WALES: along Wog Link, Coolangubra State Forest,
CGibson, 29.X.1989 (NSW); Tumblong State Forest, G.Burrows,
12.viii.l995 (NSW); 6 km WNW of Mittagong, J.Tfiompson 1656,
24.ix.1972 (NSW); track to Green Cape, M.EPhillips, 8x1961
(CANB). AUSTRALIAN CAPITAL TERRITORY: Gibraltar Creek,
N.tBurbidge 7360, 2^x.\.^962 (CANB, NSW); Black Mountain,
E.Gauba, 22j(ii.l952 (CANB). VICTORIA: Heathcote Junction
Railway Reserve, J.C.Kissane, 13.X.1984 (CANB, MEL, NSW);
turnoff to Genoa Creek, 4.6 km W of Genoa on Princes Hwy,
JHRoss 3516, 23.X.1991 (AD, BRI, CANB, HO, MEL, NSW); slopes
between Great Ocean Rd and Harvey St, Anglesea, R.V.5mith
59/247, 15.X.1959 (CANB, HO, MEL); Blacknose Point, 5 km SW
of Portland, M.D.Crisp 6925, 18.xi.l980 (CANB). TASMANIA:
Charlotte Cove, AMoscol 8660, 14.X.1984 (HO, MEL); 1.5 km S of
Cleveland on Midlands Hwy, D.LMorris 80105 (HO); Petal Point,
Cape Portland, AMosca/3207,7.X.1983 (HO, MEL); Knights Bush
near Launceston, ASimson 2264, 1889 (HO).

Flowering period: Flowers in spring.

Distribution and habitat: Occurs in south-eastern
South Australia, south-eastern Queensland, eastern
New South Wales, southern Victoria and Tasmania
(Fig. 9a). Grows in grassland, woodland and open forest.

Notes: Bossiaea prostrata is the most widespread
species in the eastern half of Australia. It can be

Muelleria

139

Thompson

distinguished from all other species in Group D by its
broad bracts and relatively proximally inserted and
caducous bracteoles, and, except for B. dasycarpa, by
its longer petioles. Two-flowered inflorescences are
more common than in other species, and occasionally
appear to be arranged in paniculate conflorescences
due to reduction or absence of subtending leaves. The
two inflorescence scales are relatively large and entire
in B. prostrata, i.e., they more closely resemble the
bract than in other species. In many species of Bossiaea
inflorescence scales are trifid rather than entire.

Although the vast majority of collections show the
faces of ovaries/valves of fruit to be glabrous, some
specimens with hairs throughout have been collected,
e.g., from Strathewen in south-central Victoria {Kilgour
466 MEL), the Grampians in south-western Victoria
{DSymon 1771 AD) and the Southern Lofty region of
South Australia (eg., Blaylock SG34 AD). The first two
of these records are in multi-piece collections and the
atypical indumentum was only present in some of the
pieces. Occasional collections with very short pedicels,
eg., Orbost, in south-eastern Victoria {Shoobridge CANB)
are thought to be aberrant development rather than
being typical of the population.

Hybridisation: A probable hybrid between
B. prostrata and B. ensata has been recorded from near
Bermagui {N.Schultz 132 CANB). It is leafy throughout
and has winged branchlets approaching the width of
those of 6 . ensata. A small sterile plant collected from
an unknown locality (Australia felix) in Victoria {F.Mueller
MEL 668111) is possibly a hybrid between Bossiaea
sericea and B. prostrata.

The Scortechinii subgroup

16. Bossiaea dasycarpa I.Thomps., sp. nov.

A. B. scortechinii F.Muell. petiolo longiore, foUolis ad
marginem minus recurvatis, leguminibus latioribus differt;
a B. prostrata R.Br. stipulis latioribus, foliolis ad marginem
recurvatis, bracteis angustioribus, bracteolis pedicello
in medio insertis persistentibus, valvis leguminis semper
hirsutis differt.

Type: Queensland. Barrett's Rd, c. 200 m N of Bruce
Highway, K.M.Sparshott 666 & K.Earnshaw, 3.xi.1995;
holotype: BRI640339; isotype: MEL 2087102.

Bossiaea prostrata var. Tuan Creek {M.S.CIemens
AQ22827).

Prostrate or decumbent subshrubs to c. 0.4 m high, with
inflorescences mostly borne on longer branchlets
rather than a regular series of short side-branchlets;
branchlets erecto-patent, mildly compressed, 0.5-0.8
mm wide, with decurrent ridges sometimes distinct,
sparsely hairy; hairs 0.3-0.5 mm long; epicuticular
wax sometimes present. Stipules triangular to narrow-
triangular, 1.5-3 mm long, with I:w ratio 2-3, erect,
herbaceous, glabrous, 1 -3-nerved; stipule-petiole angle
45-90° Leaves: petiole 1.5-4 mm long; articulation
strongly geniculate, not ridged; lamina narrow-
oblong, narrow oblong-elliptic or slightly obovate,
10-20 mm long, 3-7 mm wide, with Inv ratio mostly
1.5-3.5, flat or slightly convex each side of midrib,
mildly discolorous; base symmetrical, slightly cordate
to rounded; margin recurved to revolute, variably hairy
and tuberculate; apex truncate to subacute, sometimes
minutely acuminate with acuminate region recurved;
apiculum variably distinct, to c. 0.3 long, often brittle,
pointing down; upper surface smooth, with venation
usually raised, with gland-dotting evident, soon
glabrescent; lower surface glabrescent. Inflorescences:
axes contracted or short, sometimes multinoded,
sometimes with a leaf and stipules developed instead
of scales; inflorescences sometimes of a few flowers
in a raceme-like arrangement, with a rudimentary or
a leafy axis beyond flowers, or sometimes a solitary
axillary flower subtended by a leaf or scale arising along
a leafy branch; bract 1-1.5 mm long, 0.6-1 mm wide,
strongly convex; pedicel mostly 5-30 mm long, hairy;
bracteoles persistent or sometimes caducous at or
soon after anthesis, very narrow-elliptic, very-narrow
oblong, or lanceolate, 2-3 mm long, with l:w ratio 2-6,
mostly loosely appressed, inserted mostly in middle
third, convex, with apex nearly flat, 1-3-nerved, usually
sparsely hairy, orange-brown. Calyx A-5 mm long, hairy,
with tube c. equal to lobes; upper lobes 2-2.5 mm long,
2 mm wide, abruptly broadening at apex; lateral angle
acuminate; sinus 1.5-2 mm deep; lower lobes 2-3 mm
long, filiform distally, c. 1 mm wide, with lateral lobes flat;
standard to c. 10 mm long, slightly longer than wings
and keel, adaxially yellow with a narrow flare, abaxially
often flushed red; wings c. as long as keel, c. 2 mm
wide, mainly yellow: keel c. 3 mm wide, pale proximally,
usually red in distal third; anthers c. 0.5 mm long post¬
dehiscence; ovary hairy, 8-10-ovulate; style 3-4 mm
long. Pods: stipe 1-2 mm long; body narrow-oblong,

140

Vol 30(2) 2012

Eastern Bossiaea

20-30 mm long, 6-8 mm wide, hairy all over; hairs to c.
0.8 mm long on valves and c. 0.5 mm long on sutures;
upper margin c. 0.7 mm wide, gently convex, sometimes
minutely ridged along suture. Seeds 2.5-3 mm long, 2
mm wide; aril c. 0.8 mm long, c. 0.5 mm high, with base
c. 0.4 mm long, with lobe curving 100-150°

Selected specimens from c 12 examined: QUEENSLAND:

Gheerulla West LA, Mapleton State Forest, NW of Mapleton,
ARBean 10758, 21.ix.l996 (BRI); Childers, CH.Gittins 280,
viii.1959 (CANB); State Forest 1294 (Kullogum), c. 25.5 km
SE of Childers, K.M.Sparshott 735 & RJ.Price, 31.i.l996 (BRI);
Maryborough, M.S.C/emens, 1948 (MEL); Tuan Creek near
Maryborough, MS.CIemens, 12.X.1948 (BRI); Coast Range, 9 km
S of Biggenden, P.Young 673. ix.l983 (BRI); Mt Barney slopes,
Macpherson Range, E.F.Constable s.n., 15.xi.1952 (NSW). NEW
SOUTH WALES: E side of Emmaville Rd, 20.5 km NW of Glen
Innes, PCJobson 5154&S.AMIIs, 23.X.1997 (CANB, MEL, NSW);
Bundjalung National Park, 1.6 km S of Evans Head, R.Coveny

5116, 2.ix.l973 (NSW); c. 14 km SE of Hillgrove on Long Point
Rd, J.B.Williams, 12.X.1972 (NE); Wollomombi Falls, H.Wissman,
23.X.1963 (NSW).

Flowering period: Flowers in mid to late spring.

Distribution and habitat Occurs in south-eastern
Queensland south from the Maryborough district and in
north-eastern New South Wales as far south as Hillgrove
(Fig. 9c). Grows in woodland and grassland. Recorded
from areas of sandstone and rhyolite geology.

EtymologyiThe epithet refers to the pods which are
hairy all over (from Greek, dasys, hairy and carpos, fruit).

Notes: Bossiaea dasycarpa is similar to B. scortechinii
but has leaves with a longer petiole, less recurved
margins and a lower length to width ratio, and pods are
markedly broader. In addition bracteoles of B. dasycarpa
tend to be less linear than those of B. scortechinii, and
the keel is more strongly marked red. Bossiaea dasycarpa

Figure 9. Distributions of species in Group D. a. Bossiaea prostrata; b. B. nummularia; c. B. dasycarpa; d. B. scortechinii; e. B. obovata;

f. B. obcordata; g. B. tasmanica.

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141

Thompson

is also similar to B. prostrota but has leaves with recurved
margins, petals with smaller amounts of red markings,
narrower bracts and bracteoles, firmer herbaceous
stipules, and, in most cases, hairy pod-valves. Hairy
pod-valves have been recorded for B. prostrata in
South Australia and Victoria but not in the region of
distributional overlap with B. dasycarpa.

Specimens of B. dasycarpa have historically been
identified as either B. scortechinii or B. prostrata. Based
on annotations on a few herbarium specimens at BRI,
C.T.White planned to name B. dasycarpa as B. prostrata
var. pubicarpa; however, this name was never published.
More recently, it has been informally identified as
B. prostrata var. Tuan Creek {M.S.CIemensAQ22827).

17. Bossiaea scortechinii F.Muell., S. ScL Rec.
3(1): 1 (1883), as Scortec/i/mV

Type: [Protologue: 'On the Dumaresq River; Rev. B.
Scortechini'] Queensland. Dumaresq River, Stanthorpe,
B.Scortechini 369, date unknown; holotype: MEL 18891.

Prostrate or decumbent subshrubs to c. 0.4 m high,
with inflorescences borne on branchlets of various
lengths, but not typically on a regular series of short
side-branchlets; branchlets erecto-patent to almost
spreading, mildly compressed, 0.3-0.8 mm wide, with
decurrent ridges, moderately hairy; hairs c 0.5 mm
long; epicuticular wax sometimes developed. Stipules
narrow-triangular, 1 -3 mm long, with l:w ratio 2-3, erect
or distally recurved, herbaceous, glabrous, 1-3-nerved;
stipule-petiole angle 45-80°. Leaves: petiole 0.5-0.8
mm long; articulation strongly geniculate, not ridged;
lamina mostly narrow-oblong, sometimes oblong-
elliptic or slightly obovate, 5-15 mm long, 2-5 mm wide
(juvenile leaves broad-cuneate, c. 10 mm wide), with
l;w ratio mostly 3-5, mostly convex each side of midrib,
markedly discolorous; base rounded; margin mostly
revolute, moderately hairy, tuberculate; apex truncate
rounded or subacute, commonly strongly recurved;
apiculum to c. 0.5 mm long, generally brittle, pointing
down; upper surface smooth, with venation commonly
raised, with gland-dotting not evident, with scattered
hairs or glabrescent; lower surface with scattered hairs.
Inflorescences: axes contracted or short, sometimes
multinoded, sometimes with a leaf and stipules
developed instead of scales; inflorescences sometimes
with a few flowers in a raceme-like arrangement.

with a rudimentary or a leafy axis beyond flowers, Oj-
sometimes a solitary axillary flower subtended by a leaf
or scale arising along a leafy branch; bract 1 -2 mm long^
0.5-0.8 mm wide, moderately convex; pedicel (4-)6-2o
mm long, hairy; bracteoles persistent or sometimes
caducous soon after anthesis, narrow-oblong to linear,

1- 2.5 mm long, with l:w ratio 3-10, mostly loosely
appressed or mildly divergent, inserted mostly in middle
third, strongly convex, with apex slightly convex or flat,
sometimes filiform, 1 -nerved, sparsely hairy or glabrous,
pale yellowish or light orange-brown. Calyx 3-5 mrn
long, hairy, with tube c. equal to lobes; upper lobes 2-3
mm long, 1.5-2 mm wide, abruptly broadening at apex;
lateral angle acuminate; sinus c. 1 mm deep; lower lobes

2- 3 mm long, filiform distally, 0.7-1 mm wide, with
lateral lobes flat, with median lobe often slightly longer
than laterals; standard to c 10 mm long, slightly longer
than wings and keel, yellow, possibly without a flare;
wings c. as long as keel, c. 1.5 mm wide, yellow; keel 2-3
mm wide, pale throughout or with pink tinges apically;
anthers c. 0.3 mm long post-dehiscence; ovary hairy, c.
8-ovulate; style 2-3 mm long. Pods: stipe 1-2 mm long;
body narrow-oblong, 19-25 mm long, 4-6 mm wide,
hairy on margins and valves; hairs on valves to c. 1 mm
long, commonly appressed; hairs on sutures c. 0.5 mm
long; upper margin c. 0.7 mm wide, not ridged. Seeds
2-3 mm long, 1.5-2 mm wide; aril 0.8-1 mm long, c. 0.5
mm high, with base c. 0.4 mm long, with lobe curving
120-180° (Fig. 8c, f).

Selected specimens from c. 70 examined: QUEENSLAND;
Jibbinbar Mountain, N boundary of Sundown National Park,
PlForster 19801, 12x1996 (BRI, MEL); c. 1.5 km S of Dalveen
on Stanthorpe Rd, Darling Downs, Sl.Everist & LJ.Webb 1299,
21.xi.l946 (BRI, CANS); 11 .5 km E of Miriam Vale, EJJhompson
MIR213 & D.Baumgartner, 4.ix.1996 (BRI, MEL). NEW SOUTH
WALES: Goonoowigall State Conservation Area, c. 5 km S of
inverell, 1 km E of picnic area, LM.Cope/and4493,8.xii.2010 (BRI,
CANB, MEL, NSW); c. 15 km S of Grafton, J&P. Edwards, 2001
(NSW); Jennings, J.H.Maiden & J.LBoorman, xii.1903 (NSW);
W end of Goonoowigall Nature Reserve, 10 km S of Inverell,
GJ.White, 30.xt.l992 (NE); 'Strathbogie' 11 km W of Emmaville,
EJ.McAlister, 7.iv.1977 (NE); Demon Nature Reserve, c. 30 km SE
ofTenterfield,J.IHunfer4894,15.iii.1997 (NE).

Flowering period: Flowers mostly in spring, but also
at other times, presumably in response to rainfall.

Distribution and habitat Occurs in south-eastern
Queensland south from Miriam Vale, and in north-

142

Vol 30(2) 2012

Eastern Bossiaeo

eastern New South Wales as far south as Howell near
Inverell (Fig. 9d). Grows in sandy soils over granite in
heath, woodland and forest.

Notes: Species in the Scortechinii subgroup, and
in particular B. scortechinii, develop a number of
inflorescence variations not seen in other eastern
Australian species (see Fig. Id). These variations are
evident, however, in a number of Western Australian
species, e.g., B. loxa J.H.Ross (illustrated in Ross 2006).
Typical inflorescences terminating suppressed or very
short axes are fairly commonly seen in the Scortechinii
subgroup. However, in some plants the axis is elongated
beyond the first flower, and if two or three flowers are
present the inflorescence is raceme-like. Sometimes
these axes grow on and become leafy shoots. Sometimes
an axis appears like a normal leafy shoot, but at a node
somewhere along the axis a solitary, truly axillary flower
is borne. This flower may be subtended by a leaf or a
scale.

The Scortechinii subgroup has distinctive calyx-lobe
morphology, with lobes tapering to filiform apices. In
the upper lobes these are directed forwards. Bossiaea
scortechinii generally has the most conspicuously
filiform calyx-lobes of the subgroup, and the median
lower calyx lobe is often distinctly longer than the other
lobes. It also generally has the most slender bracteoles.

Hybridisation: A specimen from the Gara River east
of Armidale {Wallace 012/86 NSW) may be a hybrid
between B. neoanglica and B, scortechinii.

18. Bossiaea obovata I.Thomps., sp. nov.

A B. scortechinii F.Muell. foHolis obovatis, pedicellis
brevioribus, leguminibus breviorlbus, pilis leguminis
patentibus plerumque differt.

Type: New South Wales. 1 km along road to
Stanthorpe from turn-off 0.5 km N of Wilson's Downfall
on Mt Lindsay Highway, M.D.Crisp 7313 & I.RJelford,
28.ix.1974; holotype: MEL 668918; isotypes: CANB n.v.,
NSW 567111.

Prostrate or decumbent subshrubs to c. 0.2 m high, with
inflorescences borne on short to long branchlets, but
not typically on a regular series of short side-branchlets;
branchlets erecto-patent, moderately compressed,
0.3-0.8 mm wide, with well-developed decurrent ridges,
sparsely to moderately hairy; hairs 0.3-0.5 mm long;
epicuticular wax commonly developed. Stipules narrow
to very narrow-triangular, 1-2 mm long, erect or distally

recurved, herbaceous, glabrous, 1-3-nerved; stipule-
petiole angle mostly 45-80°. Leaves: petiole 0.5-1
mm long; articulation strongly geniculate, not ridged;
lamina mostly obovate or cuneate, 2-10 mm long, 1-8
mm wide, with l:w ratio mostly 1-2, flat or concave
grading to folded distally, moderately discolorous;
base symmetrical, broad-cuneate to rounded; margin
recurved to revolute, with scattered hairs and/or
tuberculate; apex truncate to subacute, or occasionally
emarginate, sometimes recurved; apiculum to 0.3 mm
long, mostly pointing down; upper surface smooth or
tuberculate, with venation raised, with gland-dotting
not evident, with scattered hairs; lower surface with
scattered hairs. Inflorescences: axes contracted or short,
or sometimes multinoded, sometimes with a leaf and
stipules developed instead of scales; inflorescences
sometimes with a few flowers in a raceme-like
arrangement, with a rudimentary or a leafy axis
beyond flowers, or sometimes a solitary axillary flower
subtended bya leaf or scale arising along a leafy branch;
bract 1-1.5 mm long, 0.3-0.6 mm wide, moderately
convex; pedicel (1-)2-5(-7) mm long, hairy; bracteoles
persistent or sometimes caducous soon after anthesis,
elliptic to narrow-oblong, 1-2 mm long, with l:w ratio
2-8, mostly loosely appressed, inserted mostly in middle
third, slightly to strongly convex, with apex flat or
slightly convex, 1 -nerved or venation obscure, glabrous,
orange-brown or reddish-brown. Calyx 3-4.5 mm long,
hairy, with tube c equal to lobes; upper lobes 2-2.5
mm long, c. 1.5 mm wide, abruptly broadening at apex;
lateral angle acute or acuminate; sinus 1.5-2 mm deep;
lower lobes 2-3 mm long, 0.8-1 mm wide, with lateral
lobes flat; standard to c. 10 mm long, slightly longer than
wings and keel, adaxially yellow with a slender flare,
abaxially sometimes flushed red; wings c. as long as
keel, 1.5-2 mm wide, yellow; keel 2.5-3 mm wide, pale
throughout or pink at apex; anthers 0.3-0.4 mm long
post-dehiscence; ovary hairy, 5- or 6-ovulate; style 3-4
mm long. Pods: stipe 1-2 mm long; body ±oblong, 12-
18 mm long, 5-7 mm wide, hairy on margins and valves,
with hairs commonly spreading; hairs on valves 1-1.5
mm long, hairs on margins 0.5-0.8 mm long; upper
margin c. 0.7 mm wide, ±flat. Seeds 2.5-3 mm long, c. 2
mm wide; aril c. 0.8 mm long, c. 0.5 mm high, with base c.
0.4 mm long, with lobe curving c. 90° (Fig. 8b).

Selected specimens from c. 80 examined: QUEENSLAND:
State Forest 639, Wrattens Forest, 22 km SSE of Kilkivan,

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143

Thompson

LPedley 5560, 15.X.1990 {BRI); Wyberba, D.Hockings, x.1963
(BRI); c. 1.6 km W of Jollys Falls, c 8 km N of Stanthorpe, LPedley
1525, 30.X.1963 (BRI); Girraween National Park, CE.Wookock,
3.xi.l983 (MEU. NEW SOUTH WALES: Burra Swamp, c 35 km
SE of Tenterfield via Spirabo Way in Forestland State Forest,
P.CJobson 5255 & S.A.Mills, 27x1997 (NSW); Wellingrove area,
M.Gray 2879, 12.iii.l954 (CANB, NSW); 11 km from Torrington
along road to Silent Grove, M.D.Crisp 7347 & I.R.Telford,
29.ix.1984 (AD, CANB, MEL); c. 1.5 km E of Cobcrofts Rd along
Mesa Management Trail, Werrikimbe National Park, LCopeland
4474, 2.xi.2010 (BRI, CANB, MEL, NE, NSW).

Flowering period: Flowers from Octoberto December,
occasionally also in autumn.

Distribution and habitat Occurs in south-eastern
Queensland, in the Stanthorpe district and also west of
Gympie, and in north-eastern New South Wales as far
south as Werrikimbe National Park, south-east ofWalcha
{Fig. 9e). Grows in sandy soils on granite, in open forest
and woodland.

Efymo/ogy: The epithet refers to the typical shape of
the leaflet lamina.

Notes: Bossiaea obovata differs from the other two
species in the Scortechinii subgroup by having obovate
leaves, short pedicels and pod-valves with spreading
hairs. Specimens of B. obovata have in the past been
assigned to B. scortechinii.

The Obcordata subgroup

19. Bossiaea tasmanica I.Thomps., nom, etstat
nov,

Bossiaea cinerea var. rigida Rodway, The Tasmanian Flora:
36(1903).

Type: [Protologue: The Rocks, near New Norfolk'.]
Tasmania. The Rocks, near New Norfolk, LRodway 168,
X.1 895; holotype: HO 1 2753.

Bossiaea obcordata sensu W.M.Curtis & D.I.Morris,
Student's FI. Tasmania, 2nd edn, 1:148 (1975).

Prostrate or decumbent shrubs to c. 0.3 m high, generally
densely and irregularly branched, with inflorescences
mostly borne on short side-branchlets; branchlets
erecto-patent to almost spreading, often recurving,
mildly compressed or c terete, 0.5-0.8 mm wide, with
decurrent ridges mostly obscure, tapering distally,
spinescent or with apex blunt, sparsely to moderately
hairy, glabrescent; hairs 0.3-0.5 mm long; epicuticular

wax commonly developed. Stipules narrow-triangular
or subulate, 0.8-1.5 mm long, erect, divergent or slightly
recurved distally, orange-brown or slightly greenish
at first, soon withering to red-brown, glabrous, faintly

1- nerved; stipule-petiole angle 0-60°. Leaves: petiole
c. 0.5 mm long; articulation strongly geniculate,
sometimes ridged; lamina elliptic to obovate, 3-7 mm
long, 2-5 mm wide, with l:w ratio mostly 1.2-2, flat or
concave grading to more strongly concave or somewhat
folded distally, mildly discolorous; base symmetrical,
rounded-cuneate; margin flat or more often recurved,
with scattered hairs, glabrescent, scarcely tuberculate;
apex subacute to truncate, commonly recurved;
apiculum to c 0.2 mm long, pointing down; upper
surface smooth, with venation sometimes slightly raised,
with gland-dotting generally evident, glabrescent; lower
surface glabrescent. Inflorescences: axes contracted;
stipules and small leaves sometimes developed instead
of scales; bract 1-1.3 mm long, strongly convex; pedicel

2- 3(-6) mm long, glabrous or sparsely hairy, glabrescent;
bracteoles commonly caducous by anthesis, narrow-
elliptic, narrow-oblong, narrow-obovate or narrow-
spathulate, 1-1.5 mm long, with l:w ratio 2-4, divergent,
inserted in proximal half, abaxial surface moderately
convex, 1-3-nerved, glabrous or sparsely hairy distally,
red-brown. Calyx 2.5-4 mm long, hairy, sometimes
sparsely so, with tube slightly longer than lobes; upper
lobes 1.5-2 mm long, 1.5-2 mm wide; lateral angle acute
or acuminate; sinus 0.5-1 mm deep; lower lobes 1.2-2.5
mm long, 0.8 mm wide, with lateral lobes flat; standard
to c. 10 mm long, c. as long as keel, adaxially yellow
with a red flare, abaxially largely brownish-red except
for pale radiating lines; wings slightly shorter than keel,
2-2.5 mm wide, purplish-brown throughout or in distal
half; keel c. 3 mm wide, pale grading to greenish-yellow,
sometimes pink-tinged at apex; anthers c. 0.6 mm long
post-dehiscence; ovary hairy, 4-ovulate; style c. 3 mm
long. Pods: stipe 1-2 mm long; body ±oblong, 15 mm
long, 6 mm wide, with hairs to 1.5 mm long on valves
and sutures; upper margin c. 0.7 mm wide, with ridge to
c. 0.3 mm high. Seeds 2.5-3 mm long, c. 1.5 mm wide; aril
c. 0.8 mm long, c. 0.8 mm high, with base c. 0.4 mm long,
with lobe curving 150-180° (Fig. 8h, j).

Selected specimens from c. 14 examined: TASMANIA: Devil
Creek headwaters, R.Barnes, 15.xi.2004 (HO); Tower Hill Rd,
M.Neyland, 12.xi.1991 (HO); 4 km S of Tunnack, B.French 628,

144

Vol 30(2) 2012

Eastern Bossiaea

7.IV.2002 (HO); Rocks near New Norfolk, LRodway, xii.1898 (HO);
Rossarden Rd, Mangara, Tleoman s.n., 8.xii.2010 (MEL).

Flowering period: Flowers in November and
December.

Distribution and habitat: Occurs in north-eastern
Tasmania near Mathinna, and in south-eastern Tasmania
south of Oatlands. Originally collected from New Norfolk
west of Hobart but not currently known from this locality
(Fig. 9g). Rare, and likely to warrant recognition as a
threatened species. Grows in loamy, gravelly or skeletal
soils derived from mudstone, in forest and woodland.

Etymology: In raising B. cinerea var, rigida to species
rank, the epithet rigido could not be used as the name
Bossiaea rigida Turcz. had already been published. The
new epithet reflects the fact that the species is endemic
to Tasmania, and B. tasmanica is in fact Tasmania's only
endemic species of Bossiaea.

Notes: Bossiaea tasmanica appears to be more closely
related than B. obcordata to other species in Group D.
It can be distinguished from B. obcordata by its more
prostrate habit, more wax-encrusted branchlets with
obscure decurrencies, blunter, branchlets that are hardly
spinous, relatively narrower leaves, narrower bracteoles,
hairy calyx and hairy pods, longer petal claws and
different petal colours.The leaves are similar in shape to
those of B. obovata of the Scortechinii subgroup.

Specimens of B. tasmanica from the type locality of
New Norfolk near Hobart in south-eastern Tasmania
have a denser indumentum than is seen in other
collections. The Rocks' as given in the protologue is
thought likely to be Derbyshire Rocks.

20. Bossiaea obcordata (Vent.) Druce, Rep, Bot.
Soc, Exch, Club Brit Isles 1916, suppl. 2:610
(1917)

Platylobium obcordatum Vent., Jard. Malmaison: subt. 31
(1803)

Type: not designated. [Cultivated in Le Jardin de
la Malmaison, France from seed collected during the
voyage of Baudin, 1802.] Holotype: G, image seen in
Geneva Herbarium Catalogue.

Platylobium microphyllum Sims, Bot. Mag. 22: 863,
pi. 863 (1805); Bossiaea microphylla (Sims) Sm., Trans.
Linn. Soc. London 9: 303 (1808). Type: not designated.
[Protologue: No information about the provenance
of seeds. Cultivated in a private garden in Berkshire,

England.] Holotype: pi. 863 in Sims, Bot. Mag. 22: 863
(1805).

Erect shrubs to c. 1 m high, with inflorescences typically
borne on a regular series of very short, side-branchlets
which in turn are produced along a regular series of
short erecto-patent side-branches; branchlets erecto-
patent, mildly compressed, 0.5-1 mm wide, with well-
developed decurrent ridges, spine-tipped, with spine
glabrous, orange-brown, sparsely to moderately hairy;
hairs to c. 0.5 mm long; epicuticular wax sometimes
developed. Stipules narrow-triangular, 1-2 mm long,
±erect, brown, sparsely hairy, glabrescent, 3-nerved;
stipule-petiole angle c. 30-60°. Leaves: petiole 0.5-1.5
mm long; articulation strongly geniculate, with ridge
absent or obscure; lamina broad-obovate, obcordate
or circular, 3-6 mm long, 2-6 mm wide, with l:w ratio
0.9-1.3, flat or gently convex each side of midrib, mostly
markedly discolorous; base symmetrical, rounded to
cuneate; margin recurved, glabrescent, ±smooth; apex
rounded, truncate or emarginate, sometimes slightly
downcurved; apiculum not or hardly developed; upper
surface smooth, with venation generally raised, with
gland-dotting sometimes evident, glabrescent; lower
surface glabrescent. Inflorescences: axes contracted
or to c. 1 mm long; bract caducous, c. 0.8 mm long,
0.6 mm wide, strongly convex; pedicel 2-4 mm long,
glabrous or sparsely hairy proximaily; bracteoles
caducous or sometimes persisting to anthesis, elliptic
to obovate, 1-1.5 mm long, with l:w ratio 1.5-2, loosely
appressed, inserted in middle or proximal thirds, slightly
to moderately convex, 3- to 5-nerved, glabrous, red-
brown. Calyx 2.5-4,5 mm long, glabrous, or occasionally
very sparsely hairy, with tube slightly to much longer
than lobes; upper lobes 1-2 mm long, 1.2-2 mm wide;
lateral angle acute; sinus 0.5-1 mm deep; lower lobes
1-1.5 mm long, c. 0.8 mm wide, with lateral lobes flat;
standard to 10 mrn long, slightly longer than wings and
keel, adaxially yellow with a red flare, and with throat
bisected, abaxially red, mainly medially; wings 0.5-1
mm shorter than keel, c. 2,5 mm wide, purplish-brown
throughout or sometimes dirty yellow distally; keel 3-4
mm wide, pinkish grading to darker red; anthers c. 0.4
mm long post-dehiscence; ovary glabrous or sparsely
hairy along upper margin, 4-ovulate; style 3-4 mm long.
Pods: stipe 2-3 mm long; body elliptic or rhomboid-

Muelleria

145

Thompson

elliptic, 10-20 mm long, 5-9 mm wide, glabrous; upper
margin c. 0.7 mm wide, with ridge to c 0.5 mm high.
Seeds 3-3.5 mm long, c. 2 mm wide; aril 1-1.5 mm
long, 1-1.5 mm high, with base c. 1 mm long, with lobe
curved 60-150° (Fig, 8g, i).

Selected specimens from c. 700 examined: QUEENSLAND:
Girraween National Park, I.RJhompson 181, 21x1995 (MEL);
Bald Rock Creek, 10 km N of Wallangarra, I.R.Telford 3180,
26.ix.1973 (CANB); Granite National Park,Wyberba,A1.S.C/emens
44729, xi.l944 (BRI). NEW SOUTH WALES: Pacific Hwy, c
20 km S of Kempsey, R.Coveny 2151, 29.ix.1969 (BRI, NSW);
Cumberland State Forest, West Pennant Hills, R.Coveny 8630,
29.X.1976 (NSW); Corner of Forest Way and Mona Vale Rd,
Belrose, R.Coveny 11902 & TJames, 20.ix.l 984 (BRI, CANB, MEL,
NSW, PERTH); c. 3 km NE of Ulan, R.Story 6818, lO.x.1959 (BRI,
CANB). VICTORIA; Brisbane Ranges National Park, c. 250 m SW
of Aeroplane Rd turnoff on Reids Rd, VStajsic 616, 26.X.1992
(MEL); W of Buchan South, A.CBeauglehole 77082, 16.ix.l984
(MEL); near Lochsport rubbish dump, The Lakes National Park,
ACBeauglehole 62840, 19j<ii.l978 (MEL).

Flowering period: Flowers from August to October.

Distribution and habitat Bossiaea obcordata occurs
in far south-eastern Queensland, eastern New South
Wales, and southern Victoria extending as far west as
the Brisbane Ranges (Fig. 9f). Grows in open forest and
woodland.

Notes: In northern New South Wales and Queensland
flowers are generally slightly larger. Bossiaea obcordata
has leaves with distinctive broad-obovate, generally
gland-dotted leaflets, moderately long petioles and a
strongly geniculate articulation.The erect, three-nerved
stipules resemble those in the Scortechinii subgroup
but are scarious and red-brown rather than green. Floral
and aril morphology and is more similar to that seen in
Group E than Group D.

The protologue of P. obcordatum in Le Jardin de la
Malmaison appears in the text associated with Plate
31, and is written in a smaller typeface. Plate 31 is an
illustration of Platylobium formosum.

Group E

Erect shrubs, sometimes moderately tall; branchelets
compressed or terete, sometimes with decurrencies,
sometimes very narrowly winged. Stipules generally
erect, sometimes with recurved, distorted, membranous
margins. Leaves with petiole mostly adaxially sulcate,
with articulation generallygeniculate; lamina sometimes

asymmetrical, generally not markedly discolorous,
with margins commonly flat and sometimes minutely
pale-rimmed, with apiculum absent or inconspicuous.
Inflorescences: axes with scales 2; bracts and bracteoles
moderately convex; pedicels relatively stout, slightly
fleshy (commonly wrinkled on drying), becoming
stouter as fruit develops; bracteoles mostly inserted
proximally. Calyx glabrous, often fleshy, drying blackish,
sometimes glaucous or glossy; upper lobes oblong
or slightly broadening from the base, mostly broader
than long, mostly not or only mildly expanded beyond
lateral angle; keel sometimes markedly elongate; wings
generally shorter than the keel; anthers relatively small.
Pods generally long-stipitate; body glabrous, with valves
mostly relatively thick and with a broad thickened upper
margin, often with seeds partitioned by spongiose
tissue internally (Fig. 10).

Group E contains nine species with seven of these
divided into two well-defined subgroups, while the
remaining two are placed in subgroups of their own.
The group extends from far north Queensland south
to Moruya in south-eastern New South Wales, and from
the coast to as much as c. 500 km inland (Fig. 11).

The Heterophylla subgroup (species 22-24) is
distinguished by compressed branchlets, short stipules,
strongly convex, basally inserted bracteoles, a calyx with
upper lobes longer than the lower lobes and moderately
expanded, and wings distinctly shorter than the keel.

The Brownii subgroup (species 25-28) is distinguished
by terete, commonly moderately hairy branchlets,
often distorted stipules with revolute membranous
margins, and leaves with an asymmetrical base. Within
the subgroup, B. carinalis and B. rupicola are clearly
distinct in having flowers with an elongate keel. The
only other species of eastern Bossiaea to have flowers
with an elongated keel is the leafless, arid-zone species
B. walkeri (Group F).

The remaining two species of Group E have a number
of peculiar features and form subgroups of their own.
Bossiaea stephensonii (21) from central-eastern New
South Wales has cladode-like branchlets, very large
erect green stipules, strongly geniculate and spurred
leaves, and an indumentum entirely of relatively long
fine hairs, while B. arenicola (29) from far northern
Queensland has bracteoles fused into a single structure,
a markedly striate calyx with distinctive triangular lobes,
and flowers with entirely yellow petals.

146

Vol 30(2) 2012

Eastern Bossiaea

Figure 10. Group E. a. 6. rhombifolia {D.E.A!brecht 5336 MEL); b. B. rupicola {J.H.Ross 3125 MEL); c. B. brownii, pod, seed among
leaves {A.N.Rodd4173 MELJ; d. R arenicola (J.CIarkson 7819 MEL); e. R brownii, stipules, base of leaflet, inflorescence {PJ.Forster
24775 MEL); f. R heterophylfa, pod-valve, inner surface {A.CBeauglehole 75827 MEL); g. comparison of upper margin of pods of
R rhombifolia (top iVoo//s MELl 528730) and R buxifolia iA.CBeauglehole68637 MEL); h. R rhombifolia, stipules {D.EAlbrecht5336
MEL); i. R concolor, stipules {I.RJhompson 486 MEL); j. R stephensonii {MJindale s.n. NSW 55359); k. R carinalis. pod (PlForster7290
MEL); I. R oligosperma, flower [I.RJhompson 1278 MEL); m. R oligosperma, leaves and immature pod (I.RJhompson 1333 MEL);
n. R arenicola, flowers U-Oorkson 7819 MEL); o. R arenicola, pod [U.Webb &J.GJracey 13610 MEL).

Scale bars: a-d = 5 mm, e, f, j, k, m-o = 2 mm, g-i = 1 mm.

Muelleria

147

.Thompson

The Stephensonii subgroup

21. Bossiaea stephensonii F.Muell., Proc. Linn.
Soc. New South Wales, ser. 2,1 (4): 1107 (1887)

Type: [Protologue:'Near Wollongong (L Stephenson,
BA)'] New South Wales. Near Wollongong, LStephenson,
x-xi.1886; holotype: MEL 20329.

Erect shrubs to c. 1 m high, with inflorescences variously
borne on longer branchlets or on a regular series of
shorter side-branchlets; branchlets erecto-patent,
strongly compressed, 1-3 mm wide, with decurrent

ridges well-developed, often narrowly winged, sparsely
to moderately hairy, generally glabrescent; hairs 1-2 mm
long; epicuticular wax not or hardly developed. Stipules
narrow-ovate to lanceolate, mostly 4-8 mm long, erect,
flat, herbaceous, glabrous, conspicuously 4-8-nerved;
stipule-petiole angle c. 45°. Leaves: petiole 1.5-3 mm
long; articulation strongly geniculate, prominently
ridged or with spur to c. 0.5 mm long; lamina narrow-
elliptic or occasionally narrow-oblong or lanceolate,
7-20(-25) mm long, 2-7 mm wide (juvenile leaves
to c. 10 mm wide), with l:w ratio mostly 2.5-5, slightly
convex each side of midrib, slightly to moderately

Key to Group E

1 Branchlets compressed, glabrous or with a sparse covering of hairs; stipules without membranous

often distorted margins.2

1: Branchlets terete, generally moderately covered by a close indumentum; stipules with

membranous margins..^...5

2 Stipules mostly > 4 mm long,green; indumentum with hairs 1-2 mm long. 2t.B.stephensonii

2: Stipules to 1.5 mm long, brown; indumentum with hairs generally < 1 mm long,

often glabrous or nearly so.3

3 All or most leaflets with l:w ratio > 2; upper calyx-lobes > the length of calyx-tube;

body of pods > 2.5 cm long.22. B. heterophylla

3: Leaflets with l:w ratio < 2; upper calyx-lobes < 1^ the length of calyx-tube; body of

pods < 2.5 cm long...4

4 Stipules to c, 0.5 mm long, c. as long as broad, with apex pointing In almost the same direction

as petiole; wing petals 10 mm long, largely brownish-red; leaves with l:w ratio mostly < 1.3.23.6. rhombifolia

4: Stipules 0.7-1.5 mm long, longer than broad, with apex pointing almost at right angles to petiole;

wing petals 6 mm long, largely yellow; leaves with l:w ratio mostly > 1.3.24. B. concolor

5 Keel 12-25 mm long, exceeding standard by several mm; persistent stamen-tube > 12 mm long;

pod-stipe 10-20 mm long.6

5: Keel to c. 10 mm long, not or hardly exceeding standard; persistent stamen-tube < 12 mm long;

pod-stipe 4-10 mm long.7

6 Leaflets with Iw ratio mostly < 4, with adaxial surface smooth, with secondary venation almost as

robust as the midrib; pedicels glabrous or with hairs generally somewhat divergent or curly.25. B. carinaiis

6: Leaflets with Iav ratio mostly > 4, with adaxial surface minutely granular, with secondary venation

markedly more slender than the midrib; pedicels with straight, appressed hairs....26. B. rupicola

7 Bracteoles fused to form a single structure; petals lacking red or purple markings; calyx

conspicuously striate; ovary hairy (northern Queensland).29. B. arenicola

7: Bracteoles free from each other; petals with red or purplish markings; calyx not conspicuously striate;

ovary glabrous (southern Queensland; New South Wales).8

8 Leaflets ovate, with base asymmetrical; body of pods 15-40 mm long, partitioned

internally (Queensland).27. B. brownii

8: Leaflets c. circular, with base c. symmetrical; body of pods 10-12 mm long,

not partitioned internally (New South Wales).28. B. oligosperma

148

Vol 30(2) 2012

Eastern Bossiaea

discolorous; base symmetrical, rounded to truncate;
margin flat to recurved, usually knobbly and pale; apex
acuminate, tapering into an apiculum; apiculum to
c. 0.1 mm long, generally pointing slightly down; upper
surface smooth or slightly tuberculate, with venation
raised, brochidodromous, with gland-dotting generally
evident, glabrescent; lower surface glabrescent.
Inflorescences: axes contracted; bract persistent, 2 mm
long, c. 1 mm wide, convex; pedicel 3-10 mm long,
glabrous; bracteoles persistent until flowering, narrow-
oblong or narrow-elliptic, 2-2.5 mm long, with l;w ratio
2.5-3, appressed, inserted near base of pedicel, strongly
convex, 3-8-nerved, glabrous, red-brown. Calyx 3-4 mm
long, glabrous, with tube equal to or longer than lobes;
upper lobes 1.2-1.8 mm long, c. 1.5 mm wide; lateral
angle acuminate; sinus 1-1.5 mm deep; lower lobes
1-1.8 mm long, 0.8-1 mm wide, with lateral lobes flat;
standard to c. 12 mm long, similar in length to or slightly
longer than wings and keel (shorter before opening),
adaxially yellow with a red flare, with throat bisected;
abaxially reddish interrupted by pale radiating nerves;
wings c. equal to keel, c. 2 mm wide, reddish proximally,
generally yellow in distal half; keel c. 3 mm wide, pink
grading to dark red; anthers c. 0.3 mm long post¬
dehiscence; ovary glabrous, 4-6-ovulate; style 3-4 mm
long. Pods: stipe 3-4 mm long; body narrow-oblong,
15-25 mm long, 6-9 mm wide, glabrous, without
spongy tissue internally; upper margin 1-1.5 mm wide,
with a ridge to c. 0.5 mm high, sometimes ridged along
suture only. Seeds 2.5-3 mm long, 1.5-1.8 mm wide; aril
0.8-1.2 mm long, c. 0.8 mm high, with base 0.7-1.2 mm
long, with lobe curving c. 90° (Fig. 1 Oj),

Selected specimens from c. 50 examined: NEW SOUTH
WALES; Port Macquarie, E.R.Brown, ii.l897 (NSW); Gan Gan
Hill, Nelsons Bay, R.Payne 2/3, viii.1993 (NSW); outskirts of
Gateshead near Newcastle, RStory 6570, 8.viii.1959 (CANB,
MEL); Morisset, J.LBoorman, x.1899 (NSW); Caley Range, Blue
Mountains National Park, W.A.CherryS76, 5.xi.2004 (NSW); Royal
National Park, just E of Engadine Railway Station, M.D.Crisp
7167, 4.X.1983 (AD, CANB, MEL); Scouters Mountain, Heathcote
National Park, R.Coveny 11607& W.Bishop, 1 .ix.l 983 (MEL, NSW).

Flowering period: Flowers in winter and spring. Most
flowers opening more or less simultaneously.

Distribution and habitat Occurs near the coast in
north-eastern and central-eastern New South Wales
from Port Macquarie in the north to Wollongong in
the south (Fig. 11a). Grows in open forest, woodland

and heathland, often in sandy soils on sandstone, but
sometimes also in clay soils.

Notes: Bossiaea stephensonii is readily identified
by its large, erect, green stipules, narrowly winged
branchlets, and long hairs. It approaches species in
Group D in terms of the relative length of lower calyx-
lobes, the relatively large, erect stipules, thin pods, long
petioles and geniculate leaflet-articulation. The bracts
and bracteoles are similar in size and colour to those of
B. prostrata.

Hybridisation: The type specimen of B. humilis Meisn.
(see under Names of uncertain application) is possibly a
hybrid involving B. stephensonii. Bossiaea obcordata is a
likely candidate as the other parent.

The Heterophylla subgroup

22. Bossiaea heterophylla Vent., Descr. PI. Nouv.
1:7,t.7(1800)

Type: not designated. [Protologue: '... originate de
Botany-Bay, introduit chez Cels en 1792, Cultivated
plant, grown, presumably, from seeds collected at
Botany Bay, New South Wales.] Holotype; t. 7 in Descr. PL
/Vouv. 1:7 (1800).

Platylobium lanceolatum Andrews, Bot Repos. 3: pi. 205
(1802); Bossiaea lanceolata (Andrews) Sm., Bot. Mag. 28:
1144, 1 .1144 (1808). Type: not designated. [Protologue:
'Our drawing was made in November 1801, at the
Nursery of Messrs. Le[e] and Kennedy, Hammersmith,
by whom it was first raised in 1792'.] Holotype: pi. 205 in
Sot. Repos. 3(1802).

Platylobium ovatum Andrews, Bot. Repos. 4: pi. 266
(1802); Bossiaea ovata (Andrews) Sm., Trans. Linn. Soc.
London 9: 303 (1808). Type: not designated. [Protologue:
'No locality information for source of seeds provided.
Cultivated at Nursery of Messr. Lee & Kennedy,
Hammersmith'.] Holotype: pi. 266 in Sot. Repos. 4: (1802).

Bossiaea heterophylla var. stenocloda Domin, Biblioth.
Bot. 22(89): 728 (1928). Type: [Protologue: 'N. S. Wales:
in der Nahe von Leura in den Blue Mts. (Domin IV.
1910)'. Near Leura, Blue Mountains, New South Wales.]
Holotype; possibly PR, n.v.

Semi-prostrate to erect shrubs to c. 2 m high, with
inflorescences typically borne on longer branchlets
rather than a regular series of short side-branchlets;
branchlets sub-erect to erecto-patent, moderately

Muelleria

149

Thompson

compressed, 1-3 mm wide, with decurrent ridges
variably well-developed, occasionally narrowly winged,
sparsely to moderately hairy, often glabrescent; hairs c.
0.2 mm long; epicuticular wax sometimes developed.
Stipules triangular, 0.5-0.8 mm long, oppressed, soon
withering to dark-brown, glabrous, with venation
indistinct; stipule-petiole angle 10-30° Leaves:
petiole 07-2.0 mm long, sulcate and hairy adaxially;
articulation strongly geniculate, often ridged; lamina
narrow oblong-elliptic, sometimes appearing almost
linear due to rolling, 4-25 mm long, 1.5-6 mm wide,
(juvenile leaves to 20 mm wide), with l:w ratio mostly
2-10, flat, incurved or involute, slightly discolorous;
base symmetrical, rounded to truncate; margin flat,
glabrous, with a smooth pale border; apex subacute
to rounded, often recurved; apiculum not developed;
upper surface smooth, with venation variably raised,
brochidodromous, with gland-dotting not evident,
glabrous; lower surface soon glabrescent Inflorescences:
axes contracted or to c. 3 mm long, moderately hairy,
occasionally with leaves instead of scales; bract 0.8-1.5
mm long, 0.8 mm wide, strongly convex; pedicel 3-8 mm
long, glabrous; bracteoles persistent, oblong to elliptic
or ovate, 0.8-2 mm long, with l:w ratio 1.5-2, generally
appressed, mostly inserted at or below mid-pedicel,
convex, with venation obscure or faintly 1-nerved,
glabrous, brown or red-brown. Calyx 3-6 mm long,
glabrous, with tube mostly slightly longer than lobes;
upper lobes 1.2-2 mm long, 1.5-3 mm wide, sometimes
slightly expanded beyond lateral angle; lateral angle
acute; sinus 1-2 mm deep; lower lobes 1-2 mm long,
c. 0.8 mm wide, with lateral lobes flat; standard to c. 15
mm long, slightly longer than keel (slightly shorter prior
to opening), adaxially yellow with a red flare, with throat
bisected; abaxially mostly flushed pink; wings 2-3 mm
shorter than keel, 1.5-2.5 mm wide, yellow throughout
or flushed pink; keel 3-4 mm wide, pink grading to
red; anthers c. 0.3 mm long post-dehiscence; ovary
glabrous, 5- or 6-ovulate; style 2-5 mm long. Pods: stipe

5- 9 mm long; body narrow-oblong, 30-45 mm long,

6- 10 mm wide, glabrous; upper margin 1.2-2 mm wide,
not ridged or with ridge to 1.5 mm high, sometimes
only finely ridged along suture; valves with transverse
venation generally obscure, with dense spongy tissue
internally. Seeds 3-4 mm long, 2-2,5 mm wide; aril 1-2
mm long, 1-1.5 mm high, with base 1-1.5 mm long,
with lobe curving 90-120° (Fig. 10f).

Selected specimens from c. 200 examined: QUEENSLAND^
near Coonarr Beach, 16 km SE of Bundaberg, L.Pedley 4425^
24Jii.l977 (BRI, CANB, NSW); Moreton Island, c. 6.5 km NE of
Tangalooma, LDurrington 1285, 14.iv.1974 (BRI); 5 km WNW
of Orchid Beach, Fraser Island, Great Sandy National Park,
P.I.Forster30253 & Gleiper, 4.ix.2004 (BRI); Pine Ridge Wildflower
Reserve, c. 8 km NNW from Southport, Blebler & P.Baxter.
30.iv.l968 (BRI). NEW SOUTH WALES: Mororo, NW of lluka.
RdFensham 4921, 20.viii.2003 (BRI); Blackheath, E.Gauba 832.
5.iii.1953 (CANB); Jervis Bay, M.E.Phillips 1109, 3.V.1961 (CANB);
Cowan to Jerusalem Bay, HSalasoo 2126, 2.iv.l 961 (NSW); Yena
Fire Trail, Captain Cook Drive, Kurnell Peninsula, Botany Bay
National Park, D.M.Crayn & R.G.Coveny 938, 28.ix.2005 (NSW);
Warrah Trig Station, Gosford, D.Gibbons, 29.iv.1987 (NSW)v
VICTORIA: N margin of Holey Plains State Park, M.G.Corrich
10034, 24.xi.1986 (MEL); near Clinton Rocks track. Captain Cook
National Park, A.CSeauglehole 33490, 20.vii.1970 (MEL); railway
line between Munro and Fernbank, c. 5 km W of Fernbank.
JJeanes 169, 29.iii.1995 (MEL); Spermwhale Head, N.A.Wakefielct
474/, 18.iv.1953 (MEL).

Flowering period: Flowers in autumn and winter
mostly.

Distribution and habitat. Occurs towards the coast
in south-eastern Queensland, south of Bundaberg,
in New South Wales, and in eastern Victoria (Fig. 11b).
Categorised as rare in Victoria (Walsh & Stajsic 2007).
Grows in heath, woodland and open forest

Notes: Bossiaea heterophylla is a widespread species
distinguished from the other two species in the
subgroup by its broader branchlets, leaflets with a
higher length to width ratio and incurved margins, and
longer calyx-lobes. In the Sydney area there is a form
with broader branchlets and leaves and larger flowers.
Domin named a new variety for a narrower-branched
and narrower-leaved form; however, the distinction
between the forms appears to be insufficiently discrete
to warrant formal taxonomic status. A third form, and
the predominant form in Queensland, differs from forms
occurring further south by having pods with a markedly
ridged upper margin, shorter keels with paler markings,
and shorter styles.

23. Bossiaea rhombifolia Sieber ex DC., Prodr.
2:117(1825)

Type: [Protologue:'Sieb! pi. exs. nov. holl. n. 354'.] New
South Wales. Location unknown. F.Sieber354; holotype:
G-DC, photo seen NSW; isotypes; MEL (4 sheets) 1528739
to 1528742), NSW 566031.

150

Vol 30(2) 2012

Eastern Bossiaea

Bossioea rotundifolia DC., Prodr. 2; 117 (1825). Type:
not designated. [Protologuei'in Nova-Hollandia orient'.]
Holotype: G-DC, image seen MEL, NSW.

Bossioea lenticularis Lodd., G.Lodd. & W.Lodd., Bot.
Cob. 13: 1238, pi. 1238 (1827), nom. ///eg. Type: not
designated. [Protologue: 'A native of New Holland: we
raised it in 1823 from seeds..']

Erect shrubs to c. 2 m high, with inflorescences typically
borne on a regular series of short to medium-length
side-branchlets; branchlets erecto-patent, moderately
compressed, 0.5-1 mm wide, with decurrent ridges
generally not well-developed, not winged, ±glabrous;
epicuticular wax generally not developed. Stipules
asymmetrically triangular, with side nearest branch
larger, 0.2-0.5 mm long, appressed, brown, glabrous,
1-nerved or venation obscure; stipule-petiole angle
10-20° Leaves: petiole 0.5-1 mm long, sulcate adaxially,
densely hairy in the sulcus; articulation strongly
geniculate, prominently ridged; lamina slightly oblate,
circular, rhomboid, or occasionally broad-obovate, 4-12
mm long, 3-12 mm wide, with l:w ratio 0.9-1.4(-1.8),
flat or gently concave, slightly or hardly discolorous;
base symmetrical, rounded or broad-cuneate; margin
flat, glabrous, smooth, with a pale rim; apex rounded,
obtuse, acute or acuminate, sometimes slightly
recurved; apiculum not developed; upper surface
smooth, with venation sometimes slightly raised, with
gland-dotting not evident, glabrous; lower surface
glabrous. Inflorescences: axes contracted; bract 0.5-0.8
mm long, c. 0.5 mm wide, strongly convex; pedicel
1-4 mm long, glabrous; bracteoles persistent, ovate,
0.8-1.5 mm long, with l:w ratio 1.5-2, loosely appressed
or slightly divergent, inserted mostly in proximal third,
especially near-basal, strongly convex, 1-nerved, or
sometimes obscurely 2- or 3-nerved, glabrous, often
pale at anthesis, becoming mid-brown. Calyx 2.7-5 mm
long, glabrous, often glaucous, with tube much longer
than lobes; upper lobes 0.6-1.5 mm long, 1.5-2.5 mm
wide, sometimes expanded beyond lateral angle by up
to c. 0.5 mm; lateral angle usually minutely acuminate;
sinus 0.5-1 mm deep; lower lobes 0.5-1 mm long,
0.7-1 mm wide, with lateral lobes flat or slightly convex,
with a medial ridge; standard to c. 12 mm long, slightly
longer than keel (but slightly shorter prior to opening).

adaxially yellow with red flare patches each side of
throat, with throat bisected, abaxially partly flushed
red; wings c. 1 mm shorter than keel, 2.5-3 mm wide,
predominantly but sometimes patchily brownish-
red; keel 3-4 mm wide, pinkish grading to dark red;
anthers c. 0.5 mm long post-dehiscence; ovary glabrous,
4-6-ovulate; style 4-5 mm long. Pods: stipe 5-7 mm
long; body oblong or oblong-elliptic, (10-)13-18 mm
long, 8-11 mm wide, glabrous, glaucous; upper margin
broadened by rounded lateral ridges; upper margin 2-3
mm wide, with sutural ridge to c. 1 mm high; valves with
transverse venation slightly raised, with spongy tissue
usually partitioning seeds internally. Seeds 3.5-4.5 mm
long, 2.5 mm wide; aril 1.5-2.2 mm long, 1-1.2 mm high,
with base 1-1.2 mm long, with lobe curving c. 135°
(Fig. 10a,g,h).

Selected specimens from c. 750 examined: QUEENSLAND:
Darling Downs: c. 6 km NNE of Wallangarra, 1.8 km SSE of Mt
Norman, M.D.Crisp 7325 & I.R.Telford, 28.ix.l 984 (BRI, CANp, MEL,
NSW);DrRobertsWaterhole,Girraween National Park, A/.G.M/a/s/r
3884, 15.ix.1994 (MEL, NSW); Jollys Falls, near The Summit,
M£.PhHlips, 20.ix.1966 (BRI). NEW SOUTH WALES: Fortis Creek,
24 km N of Grafton on road to Coaldale, D.B.Foreman 902,
23.viii.1985 (MEL, NE, NSW); Nelson Bay, 50 km NE of Newcastle,
T.J.McDonold 1916, 21.viii.1976 (BRI); Warrimoo, lower Blue
Mountains, K.A.McCo}l, 20.viii.1997 (CANB, NSW); Flagstone
Creek Rd, 2 km off the Gulf Rd, P.I.Forster 17530, 25.viii.l 995 (BRI,
MEL, NSW); N side of Timor Rd, 14.4 km W of Coonabarabran,
Warrumbungles National Park, P.CJobson 4851 & S.A.Mil!s,
3.ix.l997 (AD, CANB, MEL, NSW); 10 km W of Coonabarabran,
HStreimann 599, 6.xii.1973 (BRI, CANB); Donalds Creek,
c. 1.8 km due S of the confluence of Burra and Donald Creeks,
D.E.Albrecht5336, 28.viii.1993 (MEL).

Flowering period: Flowers in late winter and spring.

Distribution and habitat: Occurs in eastern New
South Wales north from Moruya, and in the Stanthorpe
district in far south-eastern Queensland (Fig. 11 c). Grows
in woodland and open forest.

Notes: Bossiaea rhombifolia is mostly easily
recognisable by the shape of its leaves, although
occasionally their length:width ratio becomes as high
as those of its closest relative B. concolor. The very
short stipules of B. rhombifolia are also distinctive with
only those of B. heterophyllo being similar. The stipules
are often asymmetrically triangular with the more
expanded half tending to cover the leaf-axil.

Muelleria

151

Thompson

24. Bossiaea concolor {Maiden & Betche)
I.Thomps., comb, etstat nov.

Bossiaea rhombifoHa var. concolor Maiden & Betche,
Proc. Linn. Soc. New South Wales, series 2,33:307 (1908);
B. rhombifoHa subsp. concolor (Maiden & Betche) A.T.Lee,
Contr. New South Wales Natl Herb. 4(3): 97 (1970).

Type: [Protologue: 'Araluen (J. L Boorman; August,
1907)'. Locality incorrectly cited.] New South Wales.
Narrabri, Jl.Boorman, viii.1907; holotype: NSW 43872;
isotypes: MEL 1528717, MEL 1528718.

Erect shrubs to c. 3 m high, with inflorescences typically
borne on a regular series of short to medium-length
side-branchlets; branchlets sub-erect to erecto-patent,
moderately compressed, 0.5-1 mm wide, with decurrent
ridges not well-defined, not winged, sometimes
transiently moderately hairy, glabrescent; hairs often
curled, c. 0.3 mm long; epicuticular wax generally not
developed. 5f/pu/es narrow-triangular, 0.7-1.5 mm long,
flat, erect or slightly divergent, mid-brown, glabrous,
1-nerved, stipule-petiole angle 45-90®. Leaves: petiole
0.5-1 mm long, sulcate and hairy adaxially; articulation
not or only slightly geniculate, mostly slightly ridged;
sometimes articulation obscure; lamina elliptic, oblong-
elliptic, or obovate, 3-8 mm long, 2-5 mm wide, with L*w
ratio mostly 1.2-1.8, flat or more often gently concave or
folded, nearly concolorous; base symmetrical, rounded
or broad-cuneate; margin flat, glabrous, smooth, with a
pale rim; apex rounded, subtruncate or slightly retuse,
sometimes slightly recurved; apiculum absent; upper
surface smooth, with venation mostly slightly raised,
with gland-dotting sometimes faintly evident, glabrous;
lower surface glabrous. Inflorescences: axes contracted;
bract 0.6-1 mm long, 0.4-0.5 mm wide, strongly convex;
pedicel 1-4 mm long, glabrous; bracteoles persistent,
ovate or elliptic, 0.7-1 mm long, with l:w ratio 1.5-2,
loosely appressed, inserted near base, strongly convex,
with venation obscure, glabrous, red-brown. Calyx 2-4
mm long, glabrous, with tube longer than lobes; upper
lobes 0.5-1.2 mm long, 1.2-1.8 mm wide, expanded
beyond lateral angle by up to 0.5 mm; lateral angle
acuminate; sinus c. 0.5 mm deep; lower lobes 0.5-1.5
mm long, 0.5-1 mm wide, with lateral lobes flat or more
often convex, often with a medial ridge; standard to
c. 9 mm long, slightly longer than keel, adaxially yellow
with a red flare, with throat bisected, abaxially generally
all yellow except for flare; wings 1-2 mm shorter than

keel, 1.5-2 mm wide, yellow; keel c. 3 mm wide, pale
proximally abruptly becoming red in distal half to two-
thirds; anthers c. 0.4 mm long post-dehiscence; ovary
glabrous, 4-6-ovulate; style 1.5-4 mm long. Pods: stipe
3-5 mm long; body oblong to elliptic, 15-25 mm long,
6-9 mm wide, glabrous; upper margin; upper margin
1.5-2 mm wide, broadened by rounded lateral ridges,
with or without sutural ridge to c. 1 mm high; valves
with transverse venation slightly raised, with spongy
tissue usually forming partitions internally. Seeds 3-3.5
mm long, 2-2.5 mm wide; aril 1.5-2 mm long, c. 1
mm high, with base c. 0.8 mm long, with lobe curving
100-150®(Fig. lOi).

Selected specimens from c. 100 examined: QUEENSLAND:

Burnett: 'Melrose', 15 km W of Eidsvold, AR.Bean 2294,
15.ix.l990 (BRI); near Western Creek, 'Mt Owen', c. 140 km N
of Mitchell, A.R.BeQn 25783, 27.xi.2006 (BRI, NSW); Auburn
State Forest, 18 km WSW of Mundubbera, P.I.Forster 36171,
30.viii.2009 (BRI, MEL, NSW); 0.7 km S of Booroondoo Creek,
Dingo Fence, SE of Moonie, A.ftSeon 12381, 6.ix.l997 (BRI, MEL,
NSW); Precipice National Park, P.I.Forster19683, 25.ix.1996 (AD,
BRI, MEL, NSW); S end of Fraser Island, iPLittle JT853, 9.X.1979
(BRI); 23.2 km N of Chinchilla-Wondai Rd on Burncluith Rd
c. 31.5 km NNE of Chinchilla, BJ.Lepschi & A.V.Slee 1269,
23x1993 (AD, BRI, HO, MEL); Darling Downs: Stretchworth
State Forest, SFl 55 Austin, Block 30/31, M.Bennie 156, 6.viii.2000
(BRI). NEW SOUTH WALES: Pilliga Nature Reserve, 1.4 km E of
Borah Creek Rd along Kerringle Rd, S.Donaldson 2417&B.A.Bell,
19.ix.2000 (CANB, NSW); Warialda State Forest 417, P.I.Forster
18234, 1 S.xii.l 995 (BRI, MEL, NSW); 2 km W of Coonabarabran,
H.lAston 2406, 28.viii.1983 (MEL, NSW); Narrabri, Jl.Boorman,
vi.1907 (NSW).

Flowering period: Flowers in late winter and spring.

Distribution and habitat: Occurs mostly inland from
the Great Divide, in south-eastern Queensland south
from Shoalwater Bay, and in north-eastern and central-
eastern New South Wales as far south as Mudgee
(Fig. 11 d). Grows mostly in sandy soils in woodland and
open forest.

Notes: Bossiaea concolor ls similar to B. rhombifoHa but
can be distinguished from that species by the following:
narrow-triangular stipules that form a large angle with
the petiole (compare Figure lOh & lOi), smaller flowers
and petals with different markings. In addition, leaflets
and pods on average have a higher length to width ratio
(leaflets c. 1.5 compared to c. 1.2; pods c. 3 compared
to c. 1.8), the cavities formed by spongy tissues in the

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pods are shallower, and seeds are smaller. Branchlets in
B. concolor are often slightly flexuose. The petiolule is
short with the articulation often almost in contact with
the lamina. The articulation also lacks the prominent
ridge that is typical of B. rhombifolia.

Bossiaea concolor is similar to B. nummularia in leaf-
shape and articulation and stipule morphology. Flowers
of B. concolor are relatively small in the Mundubbera
district of south-eastern Queensland, e.g., in A.R.Bean
27977 and 28063, both BRI.

The Brownii subgroup

25. Bossiaea carinatis Benth., in T.L.Mitchell,

J. Exped. Trap. Austraiia: 289 (1848)

Type: [Protologue: 'on Balmy Creek'. Balmy Creek is
in central Queensland, SE of Emerald.] Queensland.
Locality unknown (label states Sub-Tropical New
Holland), TlMtchell 275, 31.viii.1846; lectotype (here
selected): K 000278379, image seen in Kew Herbarium
Catalogue.

Residual syntypes: MEL 664781; MEL 665573;
K 000278378, image seen in Kew Herbarium Catalogue.

Erect shrubs to c. 3 m high, with inflorescences borne
variously on long branchlets or on a ±regular series of
short to medium-length side-branchlets; branchlets
erecto-patent, terete, 0.8 mm wide, without decurrent
ridges, moderately to very densely hairy; hairs mostly
c. 0.3 mm long, straight or curly, sometimes also with
short-lived spreading hairs to c. 1 mm long; epicuticular
wax sometimes developed. Stipules narrow-triangular,
2-3(-6) mm long, generally erect but sometimes
distorted, with membranous margins becoming
revolute, red-brown, hairy medially, with venation
obscure; stipule-petiole angle 80-90°. Leaves alternate;
petiole 0.5-1 mm long; articulation mostly slightly to
moderately geniculate, sometimes knobbly, sometimes
obscure; lamina narrow-ovate to lanceolate, mostly
5-25 mm long, 2-13 mm wide, with l:w ratio mostly
2-4, flat, moderately discolorous; base asymmetrical,
rounded to cordate; margin flat, glabrescent, smooth,
with a pale rim; apex acute, subacute or occasionally
rounded, minutely rounded at very tip; apiculum not
developed; upper surface smooth, with venation often
slightly raised, brochidodromous, with gland-dotting
evident, soon glabrescent; lower surface glabrescent.

Inflorescences: axes contracted; bract 1-1.5 mm long,
c. 1 mm wide, moderately convex; pedicel 3-7 mm long,
glabrous or hairy; bracteoles persistent, oblong-ovate,
0.8-1.5 mm long, with !:w ratio 1.5-2, loosely appressed,
inserted near base, often sub-opposite, deeply convex,
with venation generally obscure, glabrous or hairy
distally, red-brown. Calyx 5-9 mm long, glabrous or hairy,
glaucous or glossy, with tube equal to or slightly longer
than upper lobes; upper lobes 3-3,5 mm long, 3-4 mm
wide, expanded beyond lateral angle by 1-2 mm; lateral
angle acute or minutely acuminate; sinus 0.5-1.5 mm
deep; lower lobes 1.5-2.5 mm long, 1.2-1.5 mm wide at
base, with lateral lobes flat; standard to c. 12 mm long,
adaxially yellow, nature of flare unknown (area dries
blackish in pressed specimens), abaxially sometimes
flushed red; wings slightly longer than standard,
4-5 mm wide, yellow or red; keel 5-10 mm longer than
standard, 4-7 mm wide, pale proximally, pink to red
distally; anthers 0.8 mm long post-dehiscence; ovary
glabrous, 6-ovulate; style c. 10 mm long. Pods: stipe
10-20 mm long; body mostly toblong, 25-40 mm
long, 12-18 mm wide, glabrous; upper margin with a
thick wing 2-3 mm high abruptly broadening at the
summit to be 1.5-2 mm wide, summit flat or gently
convex; valves with transverse venation slightly raised,
with spongy internal partitions. Seeds c. 5 mm long,
2.5-3 mm wide; aril c. 2.5 mm long, c. 1.5 mm high, with
base c. 1.5 mm long, with lobe curving c. 150° (Fig. 10k).

Selected specimens from c. 700 examined: QUEENSLAND:
Coles Rd, Coominglah State Forest, W of Monto, AR.Bean
8848, 17.viii.1995 (BRI, CANB, MEL); slopes of junction ridge,
N of Marlong Arch, Mt Moffatt bp via Injune, AR.Beon 14310,
27.X.1998 (BRI); Ka Ka Mundi section, Carnarvon National Park,
SE of Mt Mooloolong, M.B.Thomas 3803, 28.viii.2008 (BRI, MEL,
NSW); Bertunya Gorge, 4 km W of Warrang homestead, White
Mountains National Park, Burke district, GAnchen 182, 17.i.l 995
(BRI, DN A, MEL); Side gorge off Rugged Gorge, c. 7 km upstream
from junction with Flinders River, White Mountains, Burke
district, MSJhomas 1631B, 13.iv.2000 (BRI); Lake Elphinstone,
ridge at southern end, P.I.Forster 7290, 26.viii.1990 (BRI, CANB,
MEL).

Flowering period: Flowers at most times of year, but
mainly in late autumn, winter, and early spring.

Distribution and habitat Occurs in eastern
Queensland between Bundaberg in the south and
Townsville in the north (Fig. 11f). Grows on sandstone in
woodland and open forest.

Muelleria

153

Thompson

Notes: Bossiaea carinalis and B. rupicola are very similar
and are not always easily distinguished. In most cases,
however, specimens can be assigned comfortably to
one or other species based on leaf length to width ratio
and differences in the indumentum on various parts.The
greater prominence of leaf venation in B. carinalis is also
a fairly reliable character as is the greater smoothness of
the adaxial surface. The calyx of fi. carinalis is sometimes
glabrous, in which case it is immediately distinguishable
from B. rupicola; however, sometimes the calyx has an
indumentum like that of B. rupicola. Fresh flowers have
not been seen by the author. The description of petal
colour varies considerably in ft carinalis, with perhaps
the majority indicating that there is some yellow evident
in standard and/or wings. In contrast, all specimen labels
indicate that these petals are red in ft rupicola.

Pods of ft carinalis and ft rupicola exhibit two
raised and broad longitudinal nerves on valves, one
approximately 2 mm in from the upper margin and the
other a similar distance in from the lower margin. These
mark the internal limit of the fusion zones betweer»
valves; these zones are relatively broad in these species.
These nerves are not evident or are poorly defined ir>
other species.

Typification: From the type material I have seen,
K 000278379 is the specimen showing the best example
of the flowers, and it was undoubtedly seen by Bentham-
I here select it as the lectotype of ft carinalis.

26. Bossiaea rupicola A.Cunn. ex Benth., FL
/li/sfro/.2:162(1864)

Type: [Protologue:'Queensland. Brisbane River; Fraser;
Mount Lindsayatan elevation of5700ft., A Cunn/ngham'.]

Figure 11 . Distributions of species in Group E. a. Bossiaea stephensonii; b. B. heterophylla; c. ft rhombifolia; d. ft concolor;
e. ft arenicola; f. ft carinalis; g. ft rupicola; h. ft brownii; i. ft oligosperma.

154

Vol 30(2) 2012

Eastern Bossiaea

New South Wales-Queensland border. Mount Lindesay,
A.Cunningham 159, vii.l 828; lectotype (here selected): K
000278380; isolectotypes: K 000278381, NSW 578605.
Images of K material seen in Kew Herbarium Catalogue.

Residual syntype: Queensland. Brisbane River, CFraser
134, 1829: K 000278383. Image seen in Kew Herbarium
Catalogue.

Shrubs or small trees to c. 4 m high, with inflorescences
borne on long branchlets or on a iregular series of short
to medium length side-branchlets; branchlets erecto-
patent, terete, c 0.8 mm wide, without decurrent ridges,
moderately hairy; hairs 0.1-0.3 mm long, typically
appressed; epicuticular wax not developed. Stipules
narrow-triangular, 2-3(-6) mm long, erect or variously
recurved or distorted, with membranous margins
becoming revolute, red-brown, hairy medially, with
venation obscure; stipule-petiole angle 80-90°. Leaves
alternate; petiole 0.5-1 mm long; articulation mostly
obscure, sometimes slightly geniculate, not ridged;
lamina oblong-lanceolate, narrow-oblong or narrow-
elliptic, 12-20 mm long, 3-6 mm wide, with l:w ratio
mostly 4-8, flat, slightly discolorous; base asymmetrical,
rounded to slightly cordate; margin flat, glabrescent,
smooth, with a rim variably distinct, often greenish; apex
acute, minutely rounded at very tip; apiculum absent-
upper surface minutely granular, with venation obscure
or slightly raised, brochidodromous, with gland-dotting
generally obscure, early glabrescent; lower surface
glabrescent. Inflorescences: axes contracted; bract
1-1.5 mm long, c. 0.8 mm wide, moderately convex;
pedicel 4-7 mm long, with appressed straight hairs;
bracteoles persistent, ovate, 1 -2 mm long, with l:w ratio
1.5-2, ±appressed, inserted near base, strongly convex,
with venation obscure, hairy, often hairy all over, pale
brown aging to red-brown. Calyx 5-7 mm long, hairy,
with tube equal to or slightly longer than upper lobes;
upper lobes 2.5-4 mm long, c. 3 mm wide, expanded
beyond lateral angle by up to c. 0.5 mm; lateral angle
acute; sinus 1 -2 mm deep; lower lobes 2-3 mm long, 1.5
mm wide, with lateral lobes flat; standard to c. 10 mm
long, adaxially yellow, with presence of flare not known
(flare zone slightly fleshy, drying brown), abaxially red;
wings slightly longer than standard, 4-5 mm wide, red;
keel 5-8 mm longer than standard, c. 6 mm wide, red

grading to darker red; anthers c. 0.8 mm long post¬
dehiscence; ovary glabrous, 6-8-ovulate; style 8-12 mm
long. Pods: stipe 10-20 mm long; body mostly loblong,
25-40 mm long, 12-18 mm wide, glabrous; upper
margin with a thick wing 2-2,5 mm high, sometimes
broadening near the summit, c. 1.5 mm wide; summit
convex or with a broad ridge c. 0.5 mm high; valves
with transverse venation slightly raised, with spongy
partitions internally. Seeds c. 5 mm long, c. 3 mm wide;
aril c. 2.2 mm long, c. 1.2 mm high, with base c. 1.2 mm
long, with lobe curving 150° (Fig. 10b).

Selected specimens from c. 80 examined: QUEENSLAND:

Kroombit Tops, State Forest 316, Parish of Clifford, SW of
Gladstone, EAEpp, vii.l 985 (BRI); near summit of Mt Walsh,
Mt Walsh National Park, Wide Bay district, J.Stanton, 2.viii.1978
(BRI); Knapps Peak, P.I.Forster 11996, 15.x.1992 (BRI); upper slope
of Mt French on road to Mt French 9.8 km W of Boonah, J.H.ffoss
3125, 21.viii.1985 (AD, BRI, CANB, HO, MEL); Kroombit State
Forest 316,51 km SSW of Calliope, N.Gibson TOI408, 15.X.1988
(BRI, MEL); Mt Gillies, 16 km SW of Rathdowney, I.RTelford
12131 & S.Donaldson, 28.vn.1996 (CANB, MEL, NE); Mt Ernest,
Mt Barney National Park, I.RTelford 12484, 7.X.2001 (NE, NSW).
NEW SOUTH WALES: Battery Hill, CJ.Dunn 122, 21 .xi.l 987 (AD,
BRI, CANB, MEL, NSW, PERTH).

Flowering period: Flowers in winter.

Distribution and habitat: Occurs mostly in the
McPherson Range in far south-eastern Queensland
and far north-eastern New South Wales, but also much
further north at KroombitTops National Park near Biloela
and Mt Walsh near Biggenden (Fig. Ilg). Grows often
amongst rocks on rhyolite, in open forest, woodland
and heathland.

Notes: Bossiaea rupicola is closely related to B. carinalis
but in most cases can be distinguished from the latter by
the higher length to width ratio of the leaflets. Further
comparisons between the two species are made in the
notes for B. carinalis.

27. Bossiaea brownii Benth., FI. Austral. 2 :163
(1864)

Type: [Protologue: 'Port Bowen, R.Brown, also
in Leichhardt's collection'.] Locality unknown, date
unknown, Lleichhardv. lectotype (here selected):
K 000278446, image seen in Kew Herbarium Catalogue;
probable isolectotype: MEL 95092.

Muelleria

155

Thompson

Residual syntypes: Queensland. Port Clinton, R.Brown,
22.vili.1802; BM 000885941, BRI 424996, CANB 278252,
CANB 371247, K 000278447, image seen in Kew
Herbarium Catalogue Brown also collected at nearby
Pine Port (Shoalwater Bay, and more specifically Akens
Island) in 1802; however, Bentham did not cite this
locality in the protologue.

Erect shrubs to c. 3 m high, with inflorescences typically
borne on a ±regular series of short to medium-length
side-branchlets; branchlets erecto-patent to almost
spreading, terete, 0.5-0.8 mm wide, without decurrent
ridges, moderately hairy with a mix of hair-types; hairs
c. 0.3 mm long, curly and loosely appressed hairs and
1-2 mm long, straight, spreading; epicuticular wax not
developed; internodes 1-3 mm long. Stipules narrow-
triangular, 1-3 mm long, mostly distorted and oriented
variously, with membranous margins becoming
revolute, red-brown, hairy, with venation obscure;
stipule-petiole angle c. 80-90° Leaves alternate; petiole
0.5-0.8 mm long; articulation mostly slightly geniculate,
not or hardly ridged, often obscured by hair; lamina
ovate to narrow-ovate, mostly 3-12 mm long, 2-8 mm
wide, with l:w ratio 1.3-1.8, flat, slightly discolorous;
base asymmetrical, cordate or with one side sometimes
truncate; margin flat, gradually glabrescent, tsmooth,
with a pale rim; apex subacute to rounded; apiculum
absent; upper surface smooth, with venation often
slightly raised, with gland-dotting not evident,
glabrescent; lower surface with loosely appressed hairs
to c. 1 mm long, occasionally glabrescent. Inflorescences:
axes contracted or rarely to 2 mm long; bract 1-2 mm
long, c. 1 mm wide, moderately convex; pedicel 0.5-6
mm long, glabrous or hairy; bracteoles persistent,
narrow-ovate, lanceolate or narrow oblong-elliptic,
1-1.5 mm long, with l:w ratio 1.5-2, divergent, inserted
in proximal third, often subopposite to alternate,
strongly convex, but with apex hardly convex, 1-nerved
or venation obscure, glabrous, red-brown. Calyx 3.5-4
mm long, glabrous, with tube longer than lobes;
upper lobes 1-1.8 mm long, c. 2 mm wide, sometimes
expanded slightly beyond lateral angle; lateral angle
acute or minutely acuminate; sinus c. 0.5-1 mm deep;
lower lobes 0.8-1.5 mm long, 1 mm wide at base, with
lateral lobes flat; standard 10 mm long, slightly longer
than keel, adaxially yellow with a red flare and with red

mark dividing throat, abaxially flushed red medially;
wings 0.5-2 mm shorter than keel, 2-3 mm wide, mostly
purplish; keel 3-4 mm wide, pink grading to dark red;
anthers c. 0.5 mm long post-dehiscence; ovary glabrous,
3- or 4-ovuIate; style 3-5 mm long. Pods: stipe 5-10 mm
long; body oblong to elliptic, 15-40 mm long, 8-15
mm wide, glabrous, glaucous or glossy; upper margin
broadened by rounded lateral ridges, 1.2-1.8 mm wide,
with medial ridge toe. 1 mm high; valves with transverse
venation raised, with spongy partitions internally. Seeds
c. 4.5 mm long, c. 2.5 mm wide; aril 2-2.5 mm long, c. 1.8
mm high, with base 2-2.5 mm long, with lobe curving
45-100° (Fig. 10c, e).

Selected specimens from c. 100 examined: QUEENSLAND;
HambowFaWs,B\ack6ownTab\e\ar\6,I.RJelford5701 &R.K.EIIyard,

11 .vi.l 977 (CANB, NSW); track 5 km E off Auburn Rd, 52 km N of
Warrego Hwy, N of Chinchilla, AN.Rodd 4173, 27.xi.1984 (BRI,
CANB, MEL, NSW);Myola, SW of Mundubbera,R/.forsfer25/77,
7x\A999 (AD, BRI, MEL); Curtis Rd, Kingaroy, AR.Bean 10645,
9.ix.1996 (BRI, CANB, MEL, NSW); Gurgeena, 6 km N of Burnettt
Hwy, NE of Mundubbera, ARBean 8807. 1 S.viii.l 995 (BRI, CANB,
MEL); camping area at SW of Lake Boemingin, Fraser Island,
N.G.Walsh 1398, 24.viii.1984 (BRI, MEL).

Flowering period: Flowers most times of the year.

Distribution and habitat: Occurs in central-eastern
and south-eastern Queensland south from the
Shoalwater Bay area, extending inland as far W as Mt
Playfair, SW of Springsure (Fig. 11h). Grows in sandy or
sandy-loam soils, commonly over sandstone, often in
gorges, in woodland and forest.

Notes: The indumentum of branchlets of B. brownii
is distinctive in that it comprises a mixture of long and
short hairs. The calyx is variously glaucous or glossy.
Leaves have a similar shape to those of B. carinalis but
are generally smaller, hairier, with a more strongly
cordate base, and with the articulation not as markedly
geniculate. Populations in the Mundubbera district have
smaller leaflets and a less hairy leaflet-articulation.

Typification: A flowering specimen from a collection
by Leichhardt and designated as K 000278446 is here
selected as the lectotype of B. brownii. It was sent by
F.Mueller to Bentham. Sheets bearing Robert Brown's
collections at Port Clinton (Port Bowen) housed at BM
(BM 000885941) and K (K 000278447) represent material
probably also seen by Bentham; however, there is no
specific indication of this.

156

Vol 30(2) 2012

Eastern Bossiaea

28. Bossiaea otigosperma A.T.Lee, Telopea 2(2):
215(1981)

Type: New South Wales.Tonalli River landing towards
Byrnes Creek, Warragamba, Al.Mitchell 434, 20.ix.1966;
holotype: NSW 285041; isotypes: BRI 278956, CANB
306843, MEL 596958.

Bossiaea sp. A sensu S.W.LJacobs & J.Pickard, Plants of
New South Wales ( 1981 ).

Erect shrubs to c. 1 m high, with inflorescences borne
typically on a tregular series of short side-branches;
branchlets erecto-patent to almost spreading, terete,
0.7-1 mm wide, without decurrent ridges, densely hairy;
hairs to c. 0.8 mm long, straight or wavy; epicuticular
wax not developed. Stipules narrow-triangular, 1-2
mm long, flat, erect or recurving, with thinner margins
generally not recurved, brown or red-brown, glabrous
except near base, 1-nerved or with venation obscure;
stipule-petiole angle 60-90°. Leaves alternate; petiole
0.3-0.8 mm long; articulation slightly geniculate, ridged,
often obscured by hair; lamina c. circular, mostly 3-6
mm long, 3-6 mm wide, with l:w ratio mostly c. 1, ±flat
or becoming concave distally, slightly discolorous; base
c symmetrical, rounded to truncate; margin flat, hairy,
±smooth, with a pale rim; apex rounded to truncate,
or abruptly recurved and acuminate; apiculum to c.
0.1 mm long; upper surface smooth, with venation
sometimes slightly raised, with gland dotting generally
obscure, generally soon glabrescent; lower surface with
somewhat persistent hairs, often moderately dense.
Inflorescences: axes contracted; bract persistent, 1 mm
long, 0.5-0.8 mm wide, slightly to moderately convex;
pedicel 1.5-3 mm long, glabrous; bracteoles persistent,
ovate, 1-1.5 mm long, with l;w ratio 1.5-2, loosely
appressed or divergent, inserted near base of pedicel,
moderately convex, 1-nerved or venation obscure,
glabrous, orange-brown. Calyx 3-4 mm long, glabrous,
sometimes slightly glaucous, with tube slightly longer
than lobes; upper lobes 1.2-1.8 mm long, 1.5-2 mm
wide, not expanded beyond lateral angle; lateral angle
acute or minutely acuminate; sinus c. 0.5 mm deep;
lower lobes 1-1.3 mm long, c. 0.8 mm wide at base, with
lateral lobes ±flat but with a distal ridge; standard to c.
10 mm long, slightly longer than keel (shorter prior to
opening); adaxially yellow with a red flare, with throat
bisected, abaxially flushed red medially; wings c. 1 mm
shorter than keel, 2.5-3 mm wide, flushed purple-brown

throughout or mainly yellow apically; keel c. 3.5-4 mm
wide, pink grading to dark red; anther c. 0.4 mm long
post-dehiscence; ovary glabrous, 2-ovulate; style 3-4
mm long. Pods: stipe 4-5 mm long; body c. elliptic,
10-12 mm long, 7-8 mm wide, glabrous; upper margin
1-1.3 mm wide, with ridge to c. 0.8 mm high; valves
with transverse venation raised, without spongy tissue
internally. Seeds 3-3.5 mm long, c. 2 mm wide; aril 1.5-2
mm long, c. 1.2 mm high, with base c. 1 mm long, with
lobe curving c. 180° (Fig. 101, m).

Selected specimens from c. 20 examined: NEW SOUTH
WALES: 2.5 km S along Claypit Rd from Windellama to Nerriga
Rd, RJohnstone2477SfA.E.Orme, 8.xm.2008 (MEL, NSW); Araluen
Valley, Mr & Mrs Shoobridge, ix.l 964 (CANB); corner of Oellen
Ford & Jacqua Rds, LRJhompson 1333, 24j<i.2010 (MEL); Tonalli
River Landing, towards Byrnes Creek, Warragamba, Al.Mitchell
277,17.xi.l964 {CANB, NSW).

Flowering period: Flowers mostly from late winter to
spring.

Distribution and habitat: Occurs in central-eastern
and south-eastern New South Wales, between
Warragamba in the north and Araluen Valley, NE
of Moruya in the south (Fig. Hi). Rare, and listed as
vulnerable under the Threatened Species Conservation
Act of New South Wales. Grows in sand and loam,
sometimes in shallow stony soils, in dry sclerophyll
forest.

Notes: Bossiaea oligosperma is characterised by a
moderately dense indumentum on branchlets and
leaves, circular leaves, and short, few-ovulate pods.
Unlike other members of the Brownii subgroup,
B. oligosperma does not develop spongiose tissue inside
pods. This is probably at least partly associated with the
fact that pods are only 2-ovulate. It appears to be most
closely related to B. brownii.

The Arenicola subgroup

29. Bossiaea arenicola J.H.Ross, Muelleria 7(3):
371 (1991)

Type: Queensland. Cook District, 4.3 km E of the
Hopevale-Starke road on the track to the Mclvor River
mouth, IR.CIarkson 5322, 14.vi.1984; holotype: MEL
665930; isotypes: MEL 1576791, NSW 787940. Also
designated as being in BRI, CANB, DNA, K, PERTH, QRS
but these n.v.

Muelleria

157

Thompson

Shrubs or trees to c. 8 m high, with inflorescences
typically borne on a regular series of side-branchlets;
branchlets erecto-patent, terete, c. 0.6 mm wide,
without decurrent ridges, moderately hairy, variably
glabrescent; hairs to c. 0.5 mm long; epicuticular wax
commonly developed. Stipules c. triangular, c. 1 mm
long, erect, brown, with the broad margins recurving,
wavy, glabrous, with venation obscure; stipule-petiole
angle 30-60°. Leaves: petiole 1-2 mm long, not sulcate
adaxially; articulation strongly geniculate, ridged;
lamina circular, broad-elliptic, somewhat rhomboidal,
ovate or broad-obovate, 8-15 mm long, 7-15 mm wide,
with l:w ratio mostly 1-1.2, flat, moderately discolorous;
base symmetrical, truncate, rounded or cuneate; margin
flat, glabrous, smooth; apex broadly rounded; apiculum
not developed; upper surface smooth, with venation
sometimes raised, soon glabrescent; lower surface
soon glabrescent. Inflorescences: axes contracted; bract
1-1.5 mm long, 0.8 mm wide, strongly convex; pedicel
1-5 mm long, glabrous or occasionally hairy; bracteoles
fused to form a single structure, persistent, 1-3 mm
long, with Iw ratio 2-3, sometimes with apex bilobed,
divergent, commonly inserted ±at base of receptacle,
sometimes c. mid-pedicel, convex, several-nerved,
glabrous, greenish-yellow or light brown. Calyx 4-5
mm long, glabrous, with conspicuously raised parallel
longitudinal venation, with tube longer than the lobes;
upper lobes c. triangular, 1.5-2 mm long, 2 mm wide;
lateral angle narrowly acute or acuminate; sinus 1-2
mm deep; lower lobes 1.3-2 mm long, 0.5-0.8 mm wide,
with lateral lobes acuminate, flat; standard to c. 13 mm
long, c. equal in length to keel, yellow; wings 1-2 mm
shorter than keel, c. 2.5 mm wide, yellow; keel c. 4 mm
wide, pale greenish-yellow; anthers c. 0.6 mm long post¬
dehiscence; ovary hairy, 2- or 3-ovulate; style 5-8 mm
long. Pods: stipe 6-8 mm long; body oblong to elliptic,
20 mm long, 9-12 mm wide, with hairs 1-1.5 mm long
on valves, c. 0.5-1 mm long on sutures, usually caducous
well before maturity; upper margin c 2 mm wide,
gently convex but hardly ridged; valves with transverse
venation markedly raised, usually with numerous bands
of papery tissue internally. Seeds 4 mm long, 3 mm wide,
brown; aril cl mmlong,c. 1 mm high, with base c. 1 mm
long, with lobe curving 90-130° (Fig. 10d, n, o).

Selected specimens from c, 20 examined: QUEENSLAND:

6.5 km W of Shelburne Bay, 5.5 km W of Messum Hill, Cape

York Peninsula, PlForster33944, 18.vi.2008 (BRI, MEL); Hopevale
Mission near Cooktown,-/.HackerBH47/, 31.vii.1983 (BRI); near
Logan Jack Creek [Jardine River National Park], H.Gitay log
2.viii.1987(BRI).

Flowering period: Flowers from April to June.

Distribution and habitat: Occurs in far north
Queensland from the tip of Cape York Peninsula south
to Cooktown (Fig. lie). Grows in sand dunes in closed
heath and shrublands.

Notes: Bossiaea arenicola is readily identified by its

uniquecalyxandbracteole morphology. It issimilarto the

Brownii subgroup in having stipules with membranous
margins. The pattern of new growth is similar to that of
Group A in which there are numerous new nodes with
fully-developed stipules but underdeveloped leaves
crowded along an axis.

Group F

Leafless shrubs, often extensively rhizomatous; cladodes
slightly to moderately compressed centrally, winged,
mostly broadly so, mostly ±glabrous, with epicuticular
wax sometimes lifting in flakes. Scales replacing stipules
at all nodes. Inflorescences: axes often with scales 4 or
more; multiple inflorescences sometimes arising from
an axil; pedicel mostly short; bracts and bracteoles
markedly convex, sometimes large, sometimes
caducous; bracteoles mostly inserted proximally. Calyx
glabrous, with upper lobes not or only slightly expanded
beyond lateral angle, sometimes triangular (Fig. 12).

Group F contains the 12 leafless species in eastern
Australia, and is divided here into four subgroups. It is
the most widespread of the groups, with the bulk of
the species occurring between far north Queensland
and Victoria. Extensions to this range are provided by
B. riparia which occurs in Tasmania, B.peninsularis which
occurs on the Eyre Peninsula in south-central South
Australia, and B. walkeri which extends across South
Australia and into Western Australia.

The Ensata subgroup (species 30-34) contains five
species with generally persistent bracts and bracteoles,
mostly only one pair of inflorescence scales, and
generally only one inflorescence per axil.

The other three subgroups differ from the Ensata
subgroup by having multiple pairs of inflorescence
scales, caducous bracts and bracteoles, and often

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Vol 30(2) 2012

Eastern Bossiaea

developing 2 or 3 inflorescences per axil. Out of the
three subgroups, the Fragrans subgroup {species 35 &
36) is probably closest to the Ensata subgroup.

The Bracteosa subgroup (species 37-40) is distinct in
having large bracts, bracteoles and distal inflorescence
■ scales, triangular upper calyx-lobes, and calyx-lobes
i that are brown and chartaceous. Prior to some very
j recent publications, Ross (2008) and McDougall (2009),
; specimens from this subgroup and the Fragrans
1 subgroup had been referred to B. bracteosa.

Bossiaea walkeri (41 ) forms a subgroup of its own and
! is distinguished from other leafless species by its striate
bracts and bracteoles, large flowers with an elongate
keel, and pods that are long, many-ovulate and with
hairs on margins.

The Ensata subgroup

30. Bossiaea ensata Sieber ex DC., Prodr. 2:117
(1825)

Type-. [Protologue; 'Sieb! pi. exs. nov.-holl. n. 434'.. in
Nova hollandia'] New South Wales. Locatiori unknown
[between Port Jackson and Blue Mountains], F.Sieber
434, date unknown; holotype: G-DC, image seen MEL,
photo NSW, fide Lee (1970); isotypes: MEL 651294, MEL
651295, MEL 651296.

Sprawling to erect rhizomatous leafless shrubs to c. 1 m
high, with cladodes to c. 20 mm wide, with inflorescences
borne on both long and shorter branchlets, sometimes
on a regular series of side-branchlets; inflorescence¬
bearing cladodes erect to erecto-patent,*l-5 mm wide.

Figure 12. Group F. a. Bossiaea vombata (J.H.Ross 3648 MEL); b. B. scolopendria {R.G.Coveny 15495 MEL); c. B. peninsularis [P.Tucker,
11 .X.2000 AD); d. B. riparia {N.G.Walsh 5530 MEL); e. B. grayi, {I.R.Telford8553 CANB); f. R milesiae, inflorescence axes with flowers in
bud (scales present but bract and bracteoles have fallen; arrow points to abscission scar of bracteole;-/.M/7es, 9.ix.1997 MEL);
g. R bombayensis, pod, scales and calyx {I.RIhompson 1327 MEL). Scale bars: a-e = 5 mm, f, g = 2 mm.

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159

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Key to Group F

1 Bracteoles generally falling before anthesis...2

1; Bracteoles generally persistent....8

2 Flowers > 15 mm long, with keel clearly longerthan standard; pods 50-60 mm long

(arid regions).™...41. B. walkeri

2: Flowers <15 mm long, with keel shorter than or c. equal to standard; pods 20-40 mm long

(not arid regions).3

3 Upper calyx-lobes somewhat quadrate, calyx-lobes not chartaceous; longest inflorescence-scale to

c 1 mm long; bracts and bracteoles < 2 mm long.4

3: Upper calyx-lobes triangular (resembling lower lobes); calyx-lobes distally brown and chartaceous;

longest inflorescence-scale > 1 mm long; bracts and bracteoles > 2 mm long.5

4 Cladodes greyish at flowering due to epicuticular wax; pedicels 1 -2.5 mm long, with bracteole

abscission scars concealed by scales.35.6. fragrans

4: Cladodes green at flowering; pedicels 2-4 mm long; bracteole abscission scars generally visible

(c. at level of scale apices or slightly more distal).36. B. milesiae

5 Largest scales of cladodes > 1 mm wide from midrib to margin, with conspicuous branching venation;

cladodes with recess at nodes up to 5 mm deep.40. B. bracteosa

5: Scales of cladodes < 1 mm wide from midrib to margin, with venation obscure; cladodes with recess at

nodes absent or < 1 mm deep.6

6 Longest cladode-scales s 2 mm long; cladodes green; plants generally infertile (Victoria only).39. B. vombata

6: Longest cladode-scales mostly < 2 mm long; cladodes greyish-green or if green then all cladodes

< 5 mm wide; seeds commonly set.7

7 Cladodes green; pods < 6 mm wide.37. B, bombayensis

7: Cladodes grey-green; pods > 6 mm wide.38. B. grayi

8 Petals without red markings except for a small flare on standard (northern Queensland).33. B. armitii

8: Petals more extensively marked than above (southern Queensland, New South Wales, Victoria).9

9 Bracteoles inserted on distal third of pedicel; cladodes with longest scales 2-2.5 mm long,

with a tuft of hairs in axils (Eyre Peninsula, South Australia).32. B. peninsularis

9: Bracteoles inserted proximal to mid-pedicel or occasionally on middle-third; cladodes with longest

scales 1 -1.5(-2) mm long, glabrous or nearly so in axils (far eastern Australia).10

10 Keel dark red, glabrous; pods < 25 mm long, s 1 mm wide, with upper margin c. 0.7 mm wide;
seeds 1.5-2 mm long; new growth with scattered hairs on faces; inflorescence scales commonly

4 or more.....34. B. riparia

10: Keel pale (or sometimes with a pink tinge), with some apical hairs; pods > 25 mm long,

7-12 mm wide, with upper margin 1-2 mm wide; seeds 3-4 mm long; new growth glabrous or

with hairs ±restricted to margins; inflorescence scales 2..11

11 Upper calyx-lobes < 2.5 mm wide; wings largely yellow (sometimes tinged red); bracteoles mostly

< 1.5 mm long; upper margin of pod 1-1.2 mm wide, with ridge angular...30. B.ensato

11: Upper calyx-lobes > 2.5 mm wide; wings purplish-brown; bracteoles mostly > 1.5 mm long;

upper margin of pod 1.5-2 mm wide, irounded. S^.B.scolopendria

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Vol 30(2) 2012

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not recessed at nodes and sometimes slightly widening,
±glabrous; marginal ridges well-defined, smooth or
with occasional tubercles; new growth ±linear in profile,
glabrous or sparsely hairy on margins; epicuticular wax
sometimes weakly developing, shed in smallish flakes,
with cladodes green at flowering. Scales 0.6-1.5 mm
long, 03-0.5 mm wide from midrib to margin, greenish
apart from midrib and margins, sometimes few-nerved
apart from midrib, glabrous or with a few hairs along
midrib, with margin glabrous. Leaves occasionally
developed and persisting towards base of stems; lamina
nearly circular, to 12 mm long. Inflorescences: axes
contracted; scales 2, 0.6-1 mm long; bract persistent,
1-1.5(-2) mm long, 0.3-0.6 mm wide, strongly convex;
pedicel 2-6 mm long, glabrous; bracteoles persistent,
ovate to narrow-ovate or narrow-oblong, 0.8-1.5(-2)
mm long, with l:w ratio 1.5-3, appressed, inserted
in proximal half, strongly convex, few-nerved, with
venation often obscure, glabrous, commonly slightly
fleshy, dark-brown. Calyx 3-4.5 mm long, glabrous,
often with dark stripes, with tube longer than lobes;
upper lobes sometimes broadening slightly from base,

1.5- 2.2 mm long, 1.5-2.4 mm wide; lateral angle acute
or acuminate; sinus 0.5-1.4 mm deep; lower lobes 1-1.8
mm long, 0.7-1 mm wide; lateral lobes flat or convex;
standard to c. 11 mm long, a few mm longer than wings
and keel, adaxially yellow with a red flare, abaxially
largely reddish, with pale radiating bands in medial
third; wings c. as long as keel, 2-2.5 mm wide, yellow,
sometimes also tinged red; keel c. 2.5 mm wide, pale
greenish-yellow, sometimes tinged pink apically, often
with hairs at distal end of fusion zone; anthers c. 0.3 mm
long post-dehiscence; ovary glabrous, 6-8-ovulate; style

2.5- 3 mm long. Pods: stipe 3-5 mm long; body narrow-
oblong, 30-40 mm long, 7-11 mm wide; upper margin
c. 1 mm wide, with ridge 0.3-0.6(-l) mm high; valves
with transverse venation hardly raised. Seeds 3-3.5 mm
long, 2-2.5 mm wide; aril 1.5-1.8 mm long, c. 1 mm high,
with base 0.8-1 mm long, with lobe curving 90-180°.

Selected specimens from c, 150 examined: QUEENSLAND:

Between Lake Benaroon and Lake Boemingen, Fraser Island,
DASmith, 15.viii.l971 (BRI); Noosa, CT.White. 21.viii.1949 (BRI);
Little Canalpin Swamp, North Stradbroke Island, KMStephens
07030713, 7.iii.2007 (BRI, NSW). NEW SOUTH WALES: C. 1.5 km
N of Lake Cathie, near Port Macquarie, D.l/erdon /57,17.viii.1969
(CANS); Anzac Pde, Matraville, R.Coveny 11290, 15.ix.l982 (MEL,
NSW); Jervis Bay, Canberra Botanic Gardens annexe, near Lake

McKenzie, CJyrrel 168, 6.X.1978 (CANB); track to Green Cape,
M£.Phillips 83, 8x1961 (CANB);Tarougra Forest Rd, 2 km E of
Bodalla along Potato Point Rd, E.Mullins 708, 6.X.1986 (CANB,
MEL, NSW). VICTORIA: entry to tip on Betka Rd, Mallacoota,
SJIorbes 2884, 14.ix.1985 (CANB, MEL); Mario Racecourse
Reserve, cl 2 km SE of Orbost, W.Hunter22, x.l 951 (MEL); c 0.5
km N of the mouth of Seal Creek, D.EAlbrecht 4844, 22.X.1991
(CANB, MEL, HO).

Flowering period: Flowers in spring.

Distribution and habitat: Occurs in near-coastal
areas of south-eastern Queensland, New South Wales
and far eastern Victoria (Fig. 13a). Categorised as rare in
Victoria (Walsh & Stajsic 2007). Grows in sandy soils in
heathland and open forest.

Notes: The bracts and bracteoles of R ensata and
R scolopendria usually appear less scarious than those
of most other species, and are sometimes slightly fleshy
medially (drying blackish). Bossiaea ensata is closest to
R scolopendria but compared to that species has smaller
and generally fewer flowers, shorter bracteoles, wing-
petals that are largely yellow, and pods that are thinner
and with the upper margin more angular. The calyx
morphology of R ensata and R scolopendria is similar to
that of species in Group E. A mutant with pure yellow
flowers has been recorded from Mororo in northern
New South Wales {Fenshom 4923 BRI).

Hybridisation: A probable hybrid between
R prostrato and B. ensata has been recorded from near
Bermagui {N.Schultz 132 CANB). It is leafy throughout
and has winged branchlets approaching the width of
those of B. ensata.

31. Bossiaea scolopendria (Andrews) Sm.,

Trans. Linn. Soc. London 9 :303 (1808)

Platylobium scolopendrium Andrews, Bot. Repos. 3: pi.
191 (1801), as scolopendrum.

Type: not designated. [Protologue: No locality or
collection details for seeds. A cultivated plant in the
Hibbertian collection'.] Holotype: pi. 191 in Bot. Repos.
3 (1801); epitype (here selected): New South Wales, St
Ives, CBurgess, 29.vii.1963: CANB 0006531.

Erect rhizomotous leafless shrubs to c. 1 m high, with
cladodes to c. 25 mm wide, with inflorescences borne
predominantly on long branchlets, occasionally on a
regular series of short side-branchlets; inflorescence-

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161

Thompson

bearing cladodes mostly sub-erect, mostly 3-12
mm wide, with no recession and sometimes a slight
widening below nodes, glabrous or sparsely hairy on
margins; marginal ridges ±smooth; new growth linear in
profile; epicuticular wax not developing, with cladodes
green at flowering. Scales 1-2 mm long, 0.4-0.6 mm
wide from midrib to margin, greenish between pale
midrib and pale margin, sometimes few-nerved apart
from midrib, with margin glabrous. Leaves occasionally
developed and persistent towards base of stems; lamina
elliptic, to c. 25 mm long. Inflorescences: axes contracted;
scales 2, 0.5-1 mm long; bract persistent. 1.5-2.5 mm
long, 0.8 mm wide, strongly convex; pedicel 2-3 mm
long, glabrous; bracteoles persistent, narrow-ovate,
narrow-elliptic or narrow-oblong, 1.5-2.5 mm long,
with I:w ratio 1.5-3, appressed, inserted in proximal half,
strongly convex, usually obscurely nerved apart from
ridged midline, glabrous, slightly fleshy, brown to dark
brown. Calyx 4-7 mm long, glabrous, often with dark
stripes, with tube equal to or slightly longer than lobes;
upper lobes broadening slightly from base, 2-3 mm
long, 2.6-3.5 mm wide, sometimes slightly expanded
beyond lateral angle; lateral angle acute or acuminate;
sinus 0.5-1 mm deep; lower lobes 1.5-2.5 mm long,
1 -1.3 mm wide; lateral lobes ±flat; standard to c. 15 mm
long, a few mm longer than wings and keel, adaxially
yellow with a red flare, abaxially reddish grading to
purplish throughout except for pale radiating bands
medially extending partway to margins; wings 3-4 mm
wide, purplish-brown throughout or at least in distal
half; keel 3-4 mm wide, pale greenish-yellow, with hairs
at distal end effusion zone; anthers c. 0.3 mm long post¬
dehiscence; ovary glabrous, 10-ovulate; style 3-3.5 mm
long. Pods: stipe 1 -4 mm long; body narrow-oblong, 30-
45 mm long, 10-12 mm wide; upper margin 1.5-2 mm
wide, with a low rounded ridge; valves with transverse
venation not or hardly raised. Seeds 3-4 mm long, 2-3
mm wide; aril 1-1.8 mm long, 1 -1.5 mm high, with base
1 -1.8 mm long, with lobe curving c. 90° (Fig. 12b).

Selected specimens from c. 150 examined: NEW SOUTH
WALES: Greenmans Valley Rd, W of Mt White, R.Coveny 11221,

10.viii.1983 (CANB, NSW); Muogamarra Nature Reserve, c. 3 km
S of the Hawkesbury River, BJlepschi 3971, 6j(ii.l998 (CANB);
21 km from Tomerong on Turpentine Rd, N side of the road,
F.W.Howe69. 12.ix.l983 (CANB, MEL, NSW); Ku-Ring-Gai Chase,
c. 25 km N of Sydney, T.R.Nlothian, 24.viii.1952 (AD); Maroota
Forest, W of Old Northern Rd, 2 km S of Forest Glen, R.G.Coveny
15495, 22.viii.1991 (AD, CANB, HO, MEL, PERTH).

Flowering period: Flowers in spring.

Distribution and habitat: Occurs in near-coastal
areas of central and southern New South Wales (Fig.
13b). Grows predominantly on sandstone, in heathland
and forest.

Notes: Bossiaea scolopendrio is similar to B. ensata
q.v. and there is some overlap in their distributions.
These two species differ from the other two species
in the Ensata subgroup by having bracteoles inserted
more proximally on the pedicel. Bossiaea scolopendrio
typically develops very long straight cladodes bearing
numerous, often 10-30, flowers. The cladode margin
of B. scolopendrio often has a distinct cellular pattern
which is discernible under moderate magnification.
The midline of cladode scales of B. scolopendrio and B.
ensata is more or less a continuation of the marginal
ridge of cladodes, and is thus more prominent than in
other leafless species. The scales of B. scolopendrio and
B. ensata are reminiscent of the stipules of R rhombifolia
and R heterophylla.

Typification: The holotype illustration shows a
good general likeness to R scolopendrio but cannot be
considered diagnostic. Furthermore, it is strange that
the ovary is drawn with hairs on margins as neither R
scolopendrio nor any other similar species such as R
ensata have been seen to develop hairs on the ovary.
An epitype, CSurgess, 29.vii.1963, CANB 0006531, to
the holotype illustration of R scolopendrio is therefore
selected here to aid the application of the name.

32. Bossiaea peninsularis I.Thomps., sp. nov.

A R ensata DC. squamis longioribus, indumento axillaris
densioribus, bracteolis pedicello in medio insertis differt.

Type: South Australia. 10 km E of Karkoo on the
south side of Mount Isabella Rd, PJuckers.n., 1 l.x.2000;
holotype: AD 110381.

Bossiaea ensata sensu J.Z.Weber, FI. S. Australia 4'^ edn
2:689(1986).

Erect rhizomatous leafless shrubs to c. 0.5 m high, with
cladodes to c. 14 mm wide, with inflorescences borne on
both long and shorter branchlets; inflorescence-bearing
cladodes erecto-patent, 2-5 mm wide, not recessed at
nodes and sometimes slightly dilating below scale,
glabrous except for hairs on margins above scale-axils;
marginal ridges mostly with occasional tubercles; new

162

Vol 30(2) 2012

Eastern Bossiaea

growth often very narrrow-elliptic in profile, moderately
hairy along margins; epicuticular wax developing, with
crusts sometimes lifting in small patches, with cladodes
grey-green at flowering. Scales 2-2.5 mm long, 0.3-0.5
mm wide from midrib to margin, brown, 3-5-nerved,
glabrous, with margin hairy or glabrous. Inflorescences:
axes contracted or to c. 2 mm long; scales 2, bract¬
like, 1-1.5 mm long, acute; bract persistent, 2-2.5 mm
long, 0.5-1 mm wide, moderately convex; pedicel 3-4
mm long, glabrous; bracteoles persistent, narrow-
ovate, 2-2.5 mm long, with l:w ratio 2-2.5, appressed
or divergent, inserted mostly in distal third, strongly
convex, several-nerved, glabrous, red-brown. Calyx
3.5-4 mm long, glabrous, with tube longer than lobes;
upper lobes 1.5-1.8 mm long, c. 1.8 mm wide; lateral
angle acute or minutely acuminate; sinus 0.5-1 mm
deep; lower lobes 1.5 mm long, 1-1.2 mm wide; lateral
lobes flat; standard to c. 10 mm long, c. 1 mm longer
than wings and keel, adaxially yellow with a red flare,
abaxially reddish to purplish with pale radiating bands
in medial third; wings slightly longer than keel, c. 2 mm
wide, pale proximally, purplish distally; keel c. 2.5 mm
wide, pale proximally, red distally, often with a few hairs
at distal end effusion zone; anthers c. 0.4 mm long post¬
dehiscence; ovary glabrous, c. 6-ovulate; style c. 2 mm
long, abruptly upeurved at base, with stigma elevated
well above anthers, conspicuously hairy. Mature pods
and seeds not seen; immature pods: stipe c. 2 mm long,
body elliptic or oblong, c. 15 mm long (Fig. 12c).

Selected specimens from c. 10 examined: SOUTH
AUSTRALIA: Hundred of Brooker, c. 70 km N of Port Lincoln,
CRAIcock 737, 28.X.1965 {AD, MEL, NSW); 33"57' S, 135"27' E,
Eyre Peninsula, S.Wr/g/if 4,26.viii.1981 (AD).

Flowering period: Flowers from August to October.

Distribution and habitat: Occurs on the Eyre
Peninsula in south-central South Australia (Fig. 13d).
Rare, and likely to warrant recognition as a threatened
species. Grows in mallee woodland.

Notes: Bossiaea peninsularis is perhaps closest to
6. ensata, in which specimens were formerly included,
and B. scolopendria, but differs from both of these by
having distally inserted, more striate and less fleshy
bracts and bracteoles, slightly longer cladode-scales
with the base more sharply delineated, and the adjacent
cladode margin moderately hairy. One collection
{P.Tucker s.n. AD) has a high proportion of 2-flowered
inflorescences.

33, Bossiaea armitii F.Muell., Fragm, 9(74): 44
(1879)

Type: [Protologue: 'Ad amnem Cave-Creek fluminis
Gilbert! in rupium fissuris; R. Daintree; ad cataractas
fluminis Herberti; W.E. Armit'.] Queensland. Herbert
River, W.E.Armlt 4, date unknown; lectotype (here
selected): MEL 651099; isolectotypes: MEL 651100, MEL
651101.

Residual syntype: Queensland. Cave Creek, Gilbert
River, R.Daintree, date unknown: MEL 651297.

Erect rhizomatous leafless shrubs to c. 3 m high, with
cladodes to c. 40 mm wide, with inflorescences borne on
both long and shorter branchlets; inflorescence-bearing
cladodes erecto-patent, 6-14 mm wide, not recessed
at nodes or recession up to 1 mm deep, sometimes
appearing recessed due to dilation of cladode below
node, glabrous except for hairs on margin immediately
above axil; marginal ridges smooth or minutely uneven;
new growth elliptic in profile; epicuticular wax often
developing, lifting in large thin flakes or sheets, with
cladodes green or greyish at flowering. Scales 1-3 mm
long, 0.3-1 mm wide from midrib to margin, brown,
several-nerved, with margin glabrous. Inflorescences:
axes contracted or compressed, to 2 mm long; scales

2- 4(-8), with cluster 1-1.5 mm long; bract persistent,
0.7-1 mm long, 0.5 mm wide, convex; pedicel 4-12
mm long, glabrous; bracteoles persistent, ovate, 0.5-1.2
mm long, with l:w ratio 1.2-2, appressed, inserted in
middle third, or rarely proximal third, strongly convex,

3- 5-nerved, glabrous, red-brown. Ca/yx 4.5-7 mm long,
glabrous, with tube longer than lobes; upper lobes 1 -2.5
mm long. 2-3.5 mm wide, sometimes slightly expanded
beyond lateral angle; lateral angle acute; sinus 0.5-2.5
mm deep; lower lobes 1-2 mm long, 0.8-1.5 mm wide;
lateral lobes flat; standard to c. 17 mm long, similar in
length to keel, adaxially yellow with a narrow red flare,
similar abaxially; wings 1-4 mm shorter than keel, c. 3
mm wide, yellow; keel c. 5 mm wide, yellow, sometimes
with hairs at distal end of fusion zone; anthers c. 1 mm
long post-dehiscence; ovary glabrous, 8-10-ovulate;
style 4-5 mm long. Pods: stipe 5-7 mm long; body
narrow-oblong, 25-30 mm long, 7-10 mm wide,
glabrous; upper margin c. 1 mm wide, minutely ridged
along suture. Seeds 3-4 mm long, 2-2.5 mm wide; aril
1.5 mm long, 0.8 mm high, with base 0.7 mm long, with
lobe curving 120-180°.

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163

Thompson

Selected specimens from c. 80 examined: NORTHERN
TERRITORY: Robinson River, LBrass, vii.1925 (BRI).
QUEENSLAND: Jowalbinna, c. 32 km SW of Laura, Garden
Creek, W,Hinton 92, iU978 (BRI); E of Baal Gammon mine, c
1 km by road N of Herberton, to Irvinebank Rd, c. 7 km W of
Herberlon, BJ.Conn&J. deCampo 1294. l.vi,1983 (BRI, CANB,
MEL, NSW); 44 km from Walsh River crossing on the Mungana-
Wrotham Park Rd, J.R.CIarkson 2806, 7.ii.1980 (DNA, NSW,
PERTH); Mount Mulligan, c. 40 km NW of Dimbulah, J.RC/ar/fson
5796, 15.iv.l985 (BRI, MEL, PERTH); 26.4 km by road towards
Forsayth from Einasleigh, Newcastle Range, K.R.McDonald3803.
6.iii.2005 (BRI, MEL); 34 km E of Forsayth towards Einasleigh,
RJ.Cumming 23617 (BRI, DNA, MEL); SW of "Silver Plains"
channel of Dinner Creek, A.Kanis2025, 19.viii.l 978 (BRI).

Flowering period: Flowers summer to autumn.

Distribution and habitat Occurs in far north
Queensland from the Iron Range south to Mount Bohle
near Charters Towers, and in the far east of the Northern
Territory where there is a single record from Robinson
River (Fig. 13c). Grows in woodland and shrubland, often
riparian and/or amongst rocks.

Notes: Bossiaea armitii is fairly uniform in floral and
fruit morphology but with some variation in cladode
shape and width. In some specimens cladodes are
distinctly elliptic and then often relatively broad,
whereas in others they are more linear as is typical of
cladodes of most species of Bossiaea. Differs from other
species in the Ensata subgroup by having longer flowers,
petals almost devoid of red markings, and wing petals
markedly shorter than the keel. It has shorter bracteoles
than B. scolopendria and B. peninsularis and they are
inserted mostly in the middle third of the pedicel rather
than proximal or distal thirds.

Typihcation: I here select MEL 651099 as the lectotype
of R armitii. It is preferred over the other type material as
it bears a pod as well as a flower.

34. Bossiaea riparia A.Cunn. ex Benth., FL
Austral, 2:166 (1864)

Type: [Protologue: 'Victoria. Maneroa, F.Mueller.
Tasmania. Derwent River, R. Brown; ... throughout ...,
1 D. Hooker. South Australia. Port Lincoln, RMuelleF.]
New South Wales. Downs of Minera, 5 miles SW from
Lake George, A.Cunningham 59, iv.l824; lectotype:
K, n.v., fide Lee (1970); isolectotype: K 000278532,
fide A.S.George, in sched. (piece a), image seen in Kew
Herbarium Catalogue.

Semi-prostrate to erect leafless shrubs to c. 1 m high, with
cladodes to c. 5 mm wide, with inflorescences borne
predominantly on side-branchlets, sometimes on a
regular series of short side-branchlets; inflorescence¬
bearing cladodes sub-erect to ereclo-patent, mostly 1-4
mm wide, with recession at nodes up to 0.4 mm deep,
occasionally with a sparse indumentum; marginal ridges
generally poorly defined, smooth or minutely uneven;
new growth linear in profile, with evenly scattered
straight hairs c. 0.3 mm long; epicuticular wax sometimes
developing, with crusts often lifting in flakes or sheets,
with cladodes green or grey-green or grey at flowering.
Sca/es 0.7-1-5 mm long, 0.3-0.5 mm wide from midrib to
margin, coppery-brown, 1 -3-nerved. Inflorescences: axes
contracted; scales 4-8, with cluster 0.5-1.5 mm long;
bract persistent until after anthesis, 0.8-1.2 mm long,
c. 0.8 mm wide, strongly convex; pedicel 1.5-5 mm long,

Figure 13. Distributions of species in the Ensata subgroup of Group F. a. Bossiaea ensata; b. B. scolopendria; c. B. armitii;

d. B. peninsularis; e. B. riparia.

164

Vol 30(2) 2012

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glabrous or hairy proximally; bracteoles persistent until
after anthesis, then variably caducous, ovate, elliptic
or obovate, 0.8-2 mm long, with l:w ratio 1-3, basally
appressed but commonly then divergent, inserted
mostly in middle third, strongly convex, 3-5-nerved or
with venation obscure, glabrous, red-brown. Calyx 2.5-4
mm long, glabrous or with scattered appressed hairs on
lobes, with tube much longer than lobes; upper lobes
0.8-1.3 mm long, 1-1.5 mm wide; lateral angle acute;
sinus c 0.5 mm deep; lower lobes 0.5-1.2 mm long, 0.6-
1 mm wide; lateral lobes flat except for a distal medial
ridge; standard to c 12 mm long, similar in length to
wings and keel, adaxially yellow with a red flare, with
throat bisected, abaxially reddish throughout or reddish
medially grading to yellow laterally: wings 2-2.5 mm
wide, yellow throughout or brownish-red throughout;
keel c. 3 mm wide, red throughout; anthers c. 0.5 mm
long post-dehiscence; ovary glabrous, 4-6-ovulate; style
3-4 mm long. Pods: stipe 2-4 mm long; body narrow-
oblong or narrow-obovate, 12-24 mm long, 4-7 mm
wide, glabrous; upper margin c. 0.7 mm wide, flat or
with fine sutural ridge to c. 0.3 mm high. Seeds 1.5-3
mm long, 1-1.8 mm wide; aril 0.7-1 mm long, 0.5-1.2
mm high, with base 0.7-1 mm long, with lobe curving
45-90° (Fig. 12d).

Selected specimens from c. 100 examined: NEW SOUTH
WALES: S side of Cave Creek, 1.5 km downstream of Blue
Waterholes, Kosciusko National Park, N.G.Wolsh 4880,9.xiU 998
(MEL); Snowy Mountains Hwy, c. 10 km E of Adaminaby,
ADuncan s.n., 3.xi.l994 (MEL, NSW); 'Mirrunga', W bank of
Murrumbidgee River, 6.5 km S of A.C.T, border, I.Crawford3175,
13.x. 1995 (CANB, NSW); Near Cooma, E.Gauba, 7.vii.l951 (AD,
CANB). A.C.T.; Pond Creek, Upper Cotter Valley, P.Gilmour 6263,
17.xi.l987 (CANB, NSW). VICTORIA: Wannon River at 4 posts
bridge, c. 22 km W of Hamilton, M.G.Corrick s.n., 1965 (MEL);
Big River, W bank, c. 300 m downstream from Fryers Creek
confluence, c. 7.5 km direct SW from Jamieson, N.G.Walsh 5771,
5.xii.2003 (CANB, MEL, NSW); Tipperary Track, E side of Sailors
Creek, SW of Hepburn, XH.Ross 3978, 11.i.l997 (CANB, MEL);
Mitta Mitta River, 3 km S by road from Mitta Mitta township,
N.G.Wolsh 6120, 14.X.2004 (MEL, NSW). TASMANIA: East
Risdon Nature Reserve, A/Wosca/ 16582, 9.X.1988 (HO, CANB);
Lake Augusta Rd, junction with old quarry track, R.Burns
147, 29.i.1990 (CANB); S side of Mersey River near Alum cliffs,
AM.Buchonan 7553, 28.xi.1985 (HO); Lake Sorell, AM.Buc/7onan
12661, 30.xi.1992 (HO); Pumphouse Point Rd, Lake St Clair,
RA.Burns, 6.ii.2003 (HO).

Flowering period: Flowers from August to December.

Distribution and habitat Occurs in far south-eastern
New South Wales, southern and eastern Victoria and
Tasmania (Fig. 13e). Categorised as rare in Victoria (Walsh
& Stajsic 2007). Grows in open forest and woodland at
low altitudes or up to c. 1000 m a.s.l.

Notes: Bossioea riparia is a variable species, with
variation in habit, width of cladodes, epicuticular wax
development, and petal markings. A form widespread
in the Southern Tablelands of New South Wales differs
reasonably consistently from other populations in being
more prostrate, having greyer, narrower cladodes, and in
having brownish rather than yellow wings. In Tasmania,
there is marked variation depending on location, with
some populations looking similar to the Victorian form
and others resembling the New South Wales Southern
Tablelands form. Much of the variation is thought to be
due to environmental factors.

Bracteoles of B. riparia are quite variable in shape;
however, a fairly consistent and distinctive feature is
the rather abrupt widening from a narrow basal portion
which has glabrous margins. Beyond this zone of
widening, the margins have the usual ciliate appearance.

Bossioea riparia also differs from other species in
the Ensata subgroup in having scattered hairs on the
faces of developing cladodes and 2 or more pairs of
inflorescence scales below a flower. In the other species
hairs are mostly restricted to margins and there is a
single pair of scales. B. bombayensis. while differing
in several important respects, shows some affinity to
B. riparia in having slender cladodes, narrow pods with
a slender wing/ridge, and hairs on faces rather than
margins of new growth. B. bombayensis shows some
affinity to B. riparia in having narrow pods with a slender
wing/ridge, and hairs on faces rather than margins of
new growth. It is also similar in cladode width.

Typification: Lee lectotypified a specimen based on
loosely attached fragments with diagnostic reproductive
material, and she gave details of the label for the
lectotype as 'Stony banks of rivulets winding through
the downs of Minera, 5 miles SW. from Lake George
'Bossiaea riparia' Cv.mss.'This specimen surprisingly is
not located in the on-line Kew Herbarium Catalogue,
and there may be some mistake. The collection details
given above are based on details presented on the label
oftheisolectotype.

Muelleria

165

Thompson

The Fragrans subgroup

35. Bossiaea fragrans K.L.McDougall, Telopea
12(3): 356 (2009)

Type: New South Wales. Central Tablelands:
Abercrombie Karst Conservation Area, K.LMcDougaH
1268, 21 .ix.2007; holotype: NSW 785656; isotypes: CANB
766110, MEL 2318267.

Erect rhizomatous leafless shrubs to c 2.5 m high, with
cladodes to c 20 mm wide, with inflorescences borne
on both long and short branchlets, but not generally on
a regular series of short side-branchlets; inflorescence¬
bearing cladodes erecto-patent, mostly 5-10 mm wide,
with recession at nodes 0.5-1 mm deep, glabrous;
marginal ridges well-defined, minutely uneven or
tuberculate; new growth somewhat elliptic in profile,
sparsely hairy on margins; epicuticular wax developing,
lifting in small flakes, with cladodes typically greyish
at flowering. Scales 1-2.5 mm long, 0.3-0.7 mm wide
from midrib to margin, coppery-brown, obscurely few-
nerved. Inflorescences: axes contracted; scales 4 or6, with
largest c. 1 mm long, 0.7-1 mm wide, with scale-cluster
1 -1.5 mm long; bract caducous or persistent, 1 -1.3 mm
long, 0.8 mm wide, moderately convex; pedicel 1-2.5
mm long, glabrous; bracteoles caducous before or after
anthesis, oblong-elliptic, 1 -1,3 mm long, with I:w ratio c.
2, divergent, inserted near base, strongly convex, with
venation obscure, glabrous, orange-brown. Calyx 3-45
mm long, glabrous, with tube much longer than lobes;
upper lobes 0.8-1 mm long, 1.2-1.6 mm wide, not or
hardly expanded beyond lateral angle; lateral angle
acute; sinus 0.5-0.8 mm deep; lower lobes 0.7-1 mm
long, not or hardly chartaceous distally; lateral lobes 0.8
mm wide, ±flat but with a medial ridge; median lobe
similar to laterals; standard to c. 12 mm long, similar
in length to wings and keel, adaxially yellow with red
marks at sides of throat, abaxially yellow, sometimes
with a red medial stripe; wings 2.5-3 mm wide, yellow;
keel c. 3 mm wide, ±red throughout; anthers c. 0.5 mm
long post-dehiscence; ovary glabrous, 5- or 6-ovuIate;
style 2.5-4 mm long. Pods: (based on McDougall 2010)
stipe 2.5-3.5 mm long; body narrow-oblong, 24-38 mm
long, 8-10 mm wide, glabrous. Seeds c. 3 mm long, c. 2
mm wide; aril not seen mature.

Selected specimens from 3 examined: NEW SOUTI-|
WALES: Abercrombie Caves, E of Grove Creek, K.LMcDougalt
999, 25.X.2001 (MEL); Abercrombie Caves, E of Grove Creek.
P.Carmen309, l.x.2006(CANB).

Flowering period: Flowers from September to
October.

Distribution and habitat Occurs in the vicinity of
Abercrombie Karst Conservation Area, south of Bathurst
in central-eastern New South Wales (Fig. 14a). Rare,
and listed as a critically endangered species under the
Threatened Species Conservation Act of New South
Wales. Grows on slate and volcanic substrates in White
Box woodland.

Notes: Bossiaea fragrans is similar to B. milesiaeq.v.The
vexillary stamen of B. fragrans is free at flowering, based
on the few samples examined.This feature has not been
recorded in other species of eastern Australian Bossiaea.

36. Bossiaea milesiae K.L.McDougall, Telopea
12(3): 356 (2009)

Type: New South Wales. South Coast, Brogo River, c.
25 km NNW of Bega (c. 1 km downstream from Brogo
Dam), KlMcDougall 1193, J.Miles & PJeuch, 12.ix.2006;
holotype: NSW 785654; isotype: CANB, MEL 2318264.

Erect rhizomatous leafless shrubs to c. 2 m high, with
cladodes to c. 10 mm wide, with inflorescences borne
mostly on short side-branchlets; inflorescence-bearing
cladodes sub-erect to erecto-patent, mostly 4-8
mm wide, with recession at nodes 0.5-0.8 mm deep,
glabrous or sometimes with hairs on margins somewhat
persistent, especially in scale-axils; marginal ridges well-
defined, minutely uneven; new growth slightly elliptic
in profile, sparsely hairy on margins; epicuticular wax
not developing, with cladodes green at flowering. Scales
1.5-2 mm long, c. 0.5 mm wide from midrib to margin,
red-brown, obscurely few-nerved. Inflorescences: axes
contracted; occasionally 2 or 3 inflorescences arising
from a single axil; scales 4-8, with largest c. 1 mm long,
0.7-1 mm wide; scale-cluster 1.5-2.2 mm long; bract
caducous or persistent until anthesis, 1.3-1.5 mm
long, c. 0.8 mm wide, moderately convex; pedicel 2-4
mm long, glabrous, becoming stout in fruit; bracteoles
caducous, often before anthesis, oblong-elliptic, 1.5-2
mm long, with l:w ratio c 2, loosely appressed, inserted

166

Vol 30(2) 2012

Eastern Bossiaea

0.5-1 mm from base, strongly convex, with venation
obscure, glabrous, orange-brown. Calyx 3.5-5 mm long,
glabrous, with tube much longer than lobes; upper
lobes 1-1.2 mm long, 1.5 mm wide, sometimes minutely
chartaceous distally; lateral angle commonly acuminate;
sinus c 1 mm deep; lower lobes 1-1.2 mm long,
minutely chartaceous distally; lateral lobes c. 0.8 mm
wide, slightly convex and ridged; median lobe similar to
laterals; standard to c. 11 mm long, similar in length to
wings and keel, adaxially yellow with a red flare, mostly
as two lateral patches, abaxially yellow; wings c. 3 mm
wide, yellow; keel c. 3.5 mm wide, red ±throughout;
anthers c. 0.5 mm long post-dehiscence; ovary glabrous,
8-ovulate; style 3.5-5 mm long. Pods: stipe 3-5 mm long;
body narrow-oblong, 25-35 mm long, 7-9 mm wide,
glabrous; upper margin 0.8-1 mm wide, with ridge to
c. 0.5 mm high. Seeds 2.5-3.5 mm long, 2-2.5 mm wide;
aril 1.5-1.8 mm long, 1-1.3 mm high, with basec. 1 mm
long, with lobe curving 90-150° (Fig. 12f).

Selected specimens from 5 examined: NEW SOUTH
WALES: lower banks of Brogo River, 0.5 km downstream from
wall of Brogo Dam, J.MHes s.n., 9.ix.1997 (MEL).

Flowering period: Flowers from August and
September.

Distribution and habitat: Occurs in the Brogo River
catchment W of Bega in far south-eastern New South
Wales (Fig. 14b). Grows in riparian open forest.

Notes: Bossiaea milesiae is very similar to B. fragrans
and there are very few collections of the two species
available for comparison. Based on the material seen,
B. milesiae differs in having cladodes always green,
longer pedicels, bracteoles inserted further from the
base of pedicel so that the abscission scars are generally
not concealed by scales (Fig. 12f), upper calyx-lobes
with the lateral angle acuminate, a longer pod-stipe,
and a standard that does not have a red stripe abaxially.

The Bracteosa subgroup

37. Bossiaea bombayensis K.L.McDougall,
Te/opea12{3):351 (2009)

Type: New South Wales. Southern Tablelands:
Shoalhaven River, Bombay, 9 km W of Braidwood,
KlMcDougaH 1325 & ClMcDougoll. IO.x.2008;
holotype: NSW 777997; isotypes: CANB, MEL 2312599.

Erect rhizomatous leafless shrubs to c. 1.5 m high with
cladodes to c. 5 mm wide, with inflorescences borne
on both long and short cladodes, but not generally on
a regular series of short side-branchlets; inflorescence¬
bearing cladodes sub-erect to erecto-patent, mostly 2-5
mm wide, not recessed at nodes or with recession to c.
0.7 mm deep, mostly soon glabrescent; marginal ridges
poorly to moderately defined, mostly minutely uneven;
new growth narrow-linear in profile, with scattered
hairs adjacent to scales, and occasional hairs elsewhere
along margins and sometimes also on faces; hairs
occasionally persisting; epicuticular wax occasionally
developing, lifting in flakes, with cladodes dark green or
grey-green. Scales l-1.5{-2) mm long, c. 0.5 mm wide
from midrib to margin, brown, with venation obscure,
with base sometimes minutely cordate. Inflorescences:
axes contracted; scales 4 or 6, with largest 1.5-2 mm
long, 1-1.5 mm wide; scale cluster 2-2.5 mm long; bract
mostly caducous at anthesis, 2-3 mm long, C..1.3 mm
wide, strongly convex; pedicel 1.5-3 mm long, glabrous,
not exceeding scale cluster or exceeding by up to 1 mm;
bracteoles caducous before anthesis, c, elliptic, 2.5-3.2
mm long, with l:w ratio 1.5-2, appressed, inserted near
base, strongly convex, with venation obscure, glabrous,
brown. Calyx 3.5-4.5 mm long, glabrous, with tube
longer than lobes; upper lobes triangular, 1-1.5 mm
long, 1-1.2 mm wide, slightly acuminate, chartaceous
distally; sinus 1-1.5 mm deep; lower lobes 1.5-2 mm
long, chartaceous distally; lateral lobes 1 mm wide, flat
except for distal median ridge; median lobe slightly
longer, wider and more convex than laterals; standard to
c.8mm long, similar in length to wings and keel, adaxially
yellow with a red flare, abaxially largely suffused red
but streakily pale medially and yellow towards lateral
margins; wings 2.5 mm wide, brownish-red proximally,
but largely yellow; keel 3.5 mm wide, grading from pale
to pink to red; anthers c. 0.6 mm long post-dehiscence;
ovary glabrous, 6-8-ovulate; style 3.5-4 mm long. Pods:
stipe 1-2.5 mm long; body narrow-oblong, 20-26 mm
long, 4-6 mm wide; upper margin 0.7-1 mm wide, flat
or with a fine sutural ridge to c. 0.3 mm high; valves
with transverse venation obscure. Seeds 2-2.5 mm long,
1.3-1.5 mm wide; aril c. 1 mm long, c. 0.5 mm high, with
base 0.6-0.8 mm long, with lobe curving c. 90° (Fig. 12g).

Selected specimens from c. 70 examined: NEW SOUTH
WALES: Shoalhaven River at Warri Bridge on Kings Hwy, c. 12 km

Muelleria

167

Thompson

direct NNW of Braidwood, I.RJhompson 1327, 24.xi.2010 (CANB,
HO, MEL); Shoalhaven River, Little Bombay, K.LMcDougall 1198,
21.ix.2006 (NSW).

Flowering period: Flowers in September and October.

Distribution and habitat Occurs north-west of
Braidwood in far south-eastern New South Wales
(Fig. 14c). Rare, and listed as vulnerable under the
Threatened Species Conservation Act of New South
Wales. Grows in riparian woodland.

Notes: Bossiaea bombayensis has the typical
inflorescence-scale, bract, bracteole and calyx features
of the Bracteosa subgroup, but has more slender
cladodes and more slender pods than the other species.

38. Bossiaea grayi K.L.McDougall, Telopea
12(3): 354 (2009)

Type: Australian Capital Territory. Murrumbidgee
River, 1 km downstream from Kambah Pool, I.RJelford
8553, ix.l980; holotype: CANB 8007070; isotypes: CANB
8007070 (sheet 2); MEL 641512, NSW 567291.

Erect rhizomatous leafless shrubs to c. 1.5 m high, with
cladodes to c 8 mm wide, with inflorescences borne on
long or short cladodes, but not generally on a regular
series of short side-branchlets; inflorescence-bearing
cladodes typically sub-erect, mostly 3-5 mm wide,
not recessed at nodes or with recession to c. 0.5 mm
deep, glabrous except for a few hairs often persisting
in axils; marginal ridges generally smooth; new growth
generally linear in outline, glabrous except for scattered
hairs on margins adjacent to scales; epicuticular wax
developing, lifting in small flakes, with cladodes grey-
green at flowering. Scales 1.3-2(-2.2) mm long, 0.5-0.8
mm wide from midrib to margin, appressed, red-brown
with pale margins, faintly 1-3-nerved. Inflorescences:
axes contracted; scales 4-8(-l 2), with largest 1.5-2 mm
long, c. 1.5 mm wide; scale-cluster 2.5-3.5 mm long;
bract variably persistent at anthesis, 3-3.5 mm long, 1.5-
1.8 mm wide, strongly convex; pedicel 2-2.5 mm long,
glabrous; bracteoles mostly caducous before anthesis,
3-3.5 mm long, with l:w ratio c. 2, appressed, inserted
near base, strongly convex, with venation obscure,
glabrous, brown. Calyx 4.5-5.5 mm long, glabrous,
with tube equal to or slightly longer than lobes; upper
lobes triangular, 1.5-2 mm long, 1-1.5 mm wide, acute,
chartaceous distally; sinus 1.5-2 mm deep; lower lobes

2-2.5 mm long, chartaceous distally; lateral lobes 1.2
mm wide, flat, with medially ridge distally; median lobe
slightly longer, broader and more convex than laterals;
standard to c. 11 mm long, similar in length to wings and
keel, adaxially yellow with a red flare, abaxially partly
flushed red with pale radiating nerves; wings c. 2 mm
wide, reddish proximally, yellow distally; keel c. 3 mm
wide, grading from pale to pink to red; anthers c. 0.5 mm
long post-dehiscence; ovary glabrous, c. 6-ovulate; style
4-5 mm long. Pods: stipe 2-4 mm long; narrow-oblong,
20-30 mm long, 6-9 mm wide; upper margin c. 0.7 mm
wide, not ridged. Seeds c. 3 mm long, c. 1.8 mm wide; aril
c. 1 mm long, c. 0.7 mm high, with base c. 0.6 mm long,
with lobe curving c. 150° (Fig. 12e).

Selected specimens from c. 8 examined: AUSTRALIAN
CAPITAL TERRITORY: Cotter Pumping Station, E.Gauba,
29.ix.1953 (CANB); Paddy's River, LPryor, 1937 (CANB);
Molonglo River, directly S of Lower Molonglo Sewage
Treatment Plant, N.Taws 310, 18.xii.l993 (CANB, MEL);
Murrumbidgee and Cotter Rivers junction, R.Cambage
2990, 5.xi.l911 (NSW). VICTORIA: Limestone Track,

c. 1.2 km from the Benambra-Wulgulmerang Rd, JAJeanes
2336, 03.ii.2010 (CANB, MEL).

Flowering period: Flowers in spring.

Distribution and habitat: Occurs in the Australian
Capital Territory along the banks of the Murrumbidgee
River and its tributaries, and in far north-eastern Victoria
(Fig. 14d). Rare, and listed as an endangered species
in the Australian Capital Territory. Grows in woodland,
with most records describing it as growing in sand or
amongst boulders on river banks.

Notes: Bossiaea grayi is very similar to B. vombata q.v.
The record given for Victoria is tentatively identified as
B. grayi based on vegetative features as it lacks flowers
and fruit. In this specimen, galls (which appear to be
replacing flowers) are formed at nodes, and these are
subtended by normal inflorescence scales that are
typical of B. grayi. The pattern of epicuticular wax on
cladodes is also typical of B. grayi.

39. Bossiaea vombata J.H.Ross, Muelleria 26:

54 (2008)

Type: Victoria. Wombat State Forest, Farm Rd, 3.9 km
from Junction of Back Settlement Rd and the Ballan-
Daylesford Rd at Korweinguboora, J.H.Ross 3647,
26.X.1 995; holotype: MEL 2043441.

168

Vol 30(2) 2012

Eastern Bossiaea

Erect rhizomatous leafless shrubs to c. 1.2 m high, with
cladodes to c 12 mm wide, with inflorescences borne on
both long and shorter branchlets, but not generally on
a regular series of short side-branchlets; inflorescence¬
bearing cladodes erecto-patent, 2-10 mm wide,
with recession at nodes 0.2-1 mm deep, mostly soon
glabrescent; marginal ridges well-defined, usually
slightly.uneven; new growth ±linear in outline, usually
transiently sparsely hairy along margins and sometimes
on the face; epicuticular wax hardly developed, not
lifting in sheets, with cladodes green at flowering. Scales
2-4 mm long, 0.7-1 mm wide from midrib to margin,
pale yellow, with venation obscure. Inflorescences: axes
contracted; occasionally 2 or 3 inflorescences arising
from a single axil; scales 4-10, with largest 1.5-2 mm
long, c. 1.5 mm wide; scale-cluster 2-3 mm long; bract
often persistent until after flowering, 2-3 mm long,
c. 1 mm wide, strongly convex; pedicel c 2 mm long,
glabrous; bracteoles caducous, narrow oblong-elliptic,
2.8-37 mm long, with l:w ratio 3-4, loosely appressed,
inserted near base, strongly convex, with venation
obscure, glabrous, brown. Calyx4-5 mm long, glabrous,
with tube equal to or longer than lobes; upper lobes
triangular, 1.7-2.2 mm long, 1.5-2 mm wide, slightly
acuminate, chartaceous distally; sinus 1.5-2 mm deep;
lower lobes 1.5-2.2 mm long, chartaceous distally;
lateral lobes 1 mm wide, flat or slightly convex distally
associated with medial ridge; median lobe slightly
longer, broader and more convex than laterals; standard
to c. 10 mm long, similar in length to wings and keel,
adaxially yellow with a red flare or flare absent, abaxially
yellow or partially suffused red; wings 2.5 mm wide,
all yellow or patchily suffused red; keel 3.5 mm wide,
pale or red throughout; anthers c. 0.6 mm long post¬
dehiscence; ovary glabrous, 4-6-ovulate; style 3-4 mm
long. Pods: stipe c. 3 mm long; body glabrous (not seen
mature). Seeds (one collection only) 2.5 mm long, 1.8
mm wide; aril 1.5 mm long, 0.8 mm high, with base 0.8
mm long, with lobe curving c. 135° (Fig. 12a).

Selected specimens from c. 10 examined: VICTORIA:
Spargo-Blakeville Rd, 120 m W of Cairns Rd intersection,
adjacent to road on N side, LMacoulay, 24.X.2009 (MEL);
Bendoc, W.Hunter, ix.l941 (MEL); Snowy River, behind WTree,
LHodge,xi.l957(MEL).

Flowering period: Flowers in spring.

Distribution and habitat: Occurs in south-central
Victoria near Daylesford and in far eastern Victoria at

Bendoc and W.Tree (Fig. 14e). Specimens from eastern
Victoria {Hodge MEL 1529684; Hunter MEL 1509814)
cannot be identified with certainty as they are sterile;
however, they are a good match for B. vombata
vegetatively. Rare, and likely to warrant recognition as a
threatened species. Grows in open forest.

Notes: When first described, only pure yellow-
flowered populations of B. vombata at the type locality
were known. Subsequently, nearby populations with
red markings have been found. Isolated plants with an
absence of red pigmentation have also been recorded
for B. ensata, B. cinerea and B. cord/gera.

Bossiaea vombata is very similar to B. gray! but
consistently has green cladodes, longer cladode-scales,
and is almost always infertile.

40. Bossiaea bracteosa F.Muell. ex Benth., FI.
Austral.!: 166 {^S64)

Type: not designated. [Protologue: 'In the Australian
Alps, on the Mitta-Mitta and Macalister rivers, at an
elevation of 3000 to 4000 ft., and on Mt Latrobe'.]
Victoria. Mitta Mitta River, F.Mueller, date not known;
lectotype: MEL 20333, fide Lee (1970).

Residual syntypes: Victoria. Macalister River, F.Mueller,
date unknown: MEL 20330, MEL 20331, MEL 20332,
MEL 20334; Victoria. Mitta Mitta River, RMueller, i.1854:
MEL 20336; Mt Latrobe (now Mt Loch), F.Mueller, date
unknown: MEL 20335.

Erect rhizomatous leafless shrubs to c. 2 m high, with
cladodes to c. 20 mm wide, with inflorescences borne
on both long and shorter branchlets; inflorescence-
bearing cladodes erecto-patent, mostly 4-12 mm wide,
with recession at nodes up to 5 mm deep, mostly soon
glabrescent; marginal ridges sharply defined, smooth;
new growth often narrowly oblong-elliptic in outline,
very sparsely hairy on margins, soon glabrescent;
epicuticular wax developing, lifting in sheets, with
cladodes green at flowering. Scales 3-5 mm long, 1-2.5
mm wide from midrib to margin, generally inset from
cladode margin, divergent, brown, multiveined, with
numerous radiating and branching veins, with base
sometimes cordate. Inflorescences: axes contracted;
occasionally 2 or 3 inflorescences arising at an axil; scales
6-10, with largest 1.5-2 mm long, 1.5-2 mm wide; scale-
cluster 2-2.5 mm long; bract caducous, 2.5-3.5 mm long.

Muelleria

169

Thompson

1-1.5 mm wide, strongly convex; pedicel 1.5-3 mm long,
glabrous, becoming stout in fruit; bracteoles caducous
before anthesis, narrow oblong-elliptic, 3.5-4.5 mm
long, with l;w ratio 2.5-3.5, loosely appressed, inserted
near base, strongly convex, but ±flat near margins, with
venation obscure or with midrib distinct, glabrous,
brown. Calyx 3.5-4.5 mm long, sometimes with a few
hairs near lobe apices, with tube slightly longer than
upper lobes; upper lobes c. triangular, 1.4-2 mm long,
1-1.3 mm wide, acute, chartaceous distally; sinus 1.5-2
mm deep; lower lobes 2-2.5 mm long, chartaceous
distally; lateral lobes 1.2 mm wide, flat, with medial ridge
distally; median lobe slightly longer, broader and more
convex than laterals; standard to c. 11 mm long, similar

in length to wings and keel, adaxially yellow with a red
flare, mostly as two lateral patches, abaxially yellow or
flushed red; wings 2.5-3 mm wide, yellow apart from
proximal red streak; keel c. 3 mm wide, red ±throughout;
anthers c. 0.5 mm long post-dehiscence; ovary glabrous,
6-8-ovulate: style 4-5 mm long. Pods: stipe 2 mm long;
body narrow-oblong, 20-32 mm long, 6-10 mm wide,
glabrous; upper margin c. 0.7 mm wide, with a ridge to
c. 0.5 mm high. Seeds 3-3.5 mm long, 1.5-2 mm wide;
aril 1-1.5 mm long, 1-1.2 mm high, with base 1-1.2 mm
long, with lobe curving 150-200°.

Selected specimens from c, 40 examined: VICTORIA: Mount
Hotham area, SJforbes 410, 20.xi.1979 (HO, MEL); Mount
Hotham development area, slope falling to Swindlers Creek,

• /

(T f

Figure 14. Distributions of species in the Fragrans, Bracteosa and Walkeri subgroups of Group F. a. Bossiaea fragrans; b. B. milesiae;
c. ft bombayensis; d. ft grayi; e. ft vombata; f. ft bracteosa; g. ft walkeri.

170

Vol 30(2) 2012

Eastern Bossiaea

N.G.Walsh 542, 27.xi.l 980 (AD, MEL); 2.9 km along road to Dargo
from Hotham Rd [Great Alpine Rd], LRJhompson s.n., 12.ii.1994
(MEL); Dargo High Plains, Long SpurTrack, c 2 km S from 4WD
track to Mayfield, N.G.Walsh 5716,2.12003 (CANB, MEL).

Flowering period: Flowers from November to early
January.

Distribution and habitat Occurs in the Dargo High
Plains and near Mt Hotham and near the headwaters
of the Macalister River in eastern Victoria (Fig. 14f).
Categorised as rare in Victoria (Walsh & Stajsic 2007).
Grows at 1000-1600 m above sea level in shallow soils
in Snow Gum woodland.

Notes: Bossiaea bracteosa is one of a group of four
species, the others being B. bomboyensis, B. grayi and
B. vombata, which have triangular upper calyx-lobes,
calyx-lobes distally chartaceous, and large, caducous
bracts and bracteoles. The scales of cladodes of
B. bracteosa, in addition to being larger than in other
leafless species, have much more conspicuous reticulate
venation, are sometimes cordate-based, are inserted
more deeply within the nodal recesses and are more
divergent. The scales are also unique in that their
insertion is commonly inset relative to the cladode
margin. The bract and bracteoles have recurved,
membranous margins.

The recognition by McDougall (2009) of four new
species of Bossiaea from material previously placed
in B. bracteosa has considerably narrowed the current
circumscription of R bracteosa.

The Walkeri subgroup

41. Bossiaea walkeri F.MuelL, Fragm. 2(15): 120
(1861)

Type: [Protologue: 'In pinetis montium Peel-Range
inter flumina Lachlan et Murrumbidgee. Alex. Walker'.
The Peel Range is now known as the Cocoparra Range.]
New South Wales. Cocoparra Range, A.Walker s.n.,
10.xi.1860; lectotype: MEL 20337, fide Ross (2006).

Erect leafless shrubs to c. 3 m high, with cladodes to
c. 7 mm wide, with inflorescences typically borne on
short side-branchlets; inflorescence-bearing cladodes
erecto-patent, mostly 2-6 mm wide, with recession at
nodes 0.3-1 mm deep, glabrescent; marginal ridges
well-defined, generally smooth; new growth linear in
profile, sparsely or occasionally densely hairy all over,
glabrescent; epicuticular wax generally developed, with

crusts lifting in sheets, grey-green at flowering. Juven/7e
leaves sometimes present near base of stems, with
lamina broad-elliptic, to c. 16 mm long. Scales 1.5-2.5
mm long, 0.6-1 mm wide from midrib to margin, dark
brown, with multiple veins faintly evident. Inflorescences:
axes contracted: scales 4-10, with cluster 1.5-2 mm long;
bract caducous before anthesis, 3 mm long, 1.5-2 mm
wide, convex; pedicel 2-7 mm long, glabrous or hairy
proximally; bracteoles caducous before anthesis, elliptic,
c. 3 mm long, with l:w ratio 2-2.5, divergent, inserted in
proximal third, convex, with margins outcurved, many-
nerved, glabrous, orange-brown. Calyx 7-^0 mm long,
with tube slightly longer than upper lobes; upper lobes
4-5 mm long, 3-4 mm wide, mostly expanded beyond
lateral angle by 0.5-1 mm; lateral angle acute; sinus 0.5-
1 mm deep; lower lobes 3-4 mm long; lateral lobes 1.5
mm wide, flat; median lobe similar to laterals; standard
15-20 mm long, adaxially yellow with a red flare,
abaxially suffused red; wings 1 or 2 mm shorter than
keel, 3.5-4 mm wide, brownish-red; keel 3-5 mm longer
than standard, 5-6 mm wide, red; anthers c. 0.7 mm
long post-dehiscence; ovary hairy, c. 20-ovulate; style
c. 8 mm long. Pods: stipe 3-4 mm long; body narrow-
oblong, 50-60 mm long, 7-12 mm wide; upper margin
c. 0.8 mm wide, with a ridge to c. 0.5 mm high. Seeds
3_4 mm long, c. 2 mm wide; aril 2-2.5 mm long, 1.5 mm
high, with base c. 1 mm long, with lobe curving 180°.

Selected specimens from c. 200 examined: SOUTH
AUSTRALIA: Yellabinna Regional Reserve, Nullarbor Region,
FJ.Bodman 12224, 11.viii.2006 (AD, CANB, MEL); 37.6 km
from Yardea homestead on road to Minnipa, J.D.Briggs 1137,
7.ix.l983 (AD, CANB, MEL. NSW); Chowilla Station, c. 20 km
NE of Renmark, E.Robertson, 26.viii.1974 (AD, BRI, CANB). NEW
SOUTH WALES: Mandleman station on Cobb Hwy to Mildura
Rd, R.CWeston 142, 21.viii.1988 (AD, BRI, CANB); Bundure
Station, N of Mt Hope, P.Mortensz 158, 22.V.1969 (CANB, NSW).
VICTORIA: Wyperfeld National Park, c. 1 km E of Cambacanya
clearing, ACBeauglehole 28834, 2.X.1968 (AD, MEL); Boundary
Bend, c. 30 km N of Kenley, ABegg,29.vii.l962 (AD, MEL).

Flowering period: Flowers in winter and spring.

Distribution and habitat: Occurs in arid regions of
southern Western Australia, South Australia, western
New South Wales and far north-western Victoria (Fig.
14g). Grows mostly in mallee woodland, often in red
sandy soils.

Notes: Bossiaea walkeri is distinguished from other
leafless species in eastern Australia by large flowers with
an elongate keel, thin, outrolling cladode-scale margins.

Muelleria

171

Thompson

hairy pod-margins, and seeds with a knobbly aril. Bracts
and bracteoles are distinctively striate, and margins of
these and inflorescence scales are relatively long-ciliate.
It is the only species of Bossiaea in eastern Australia to
occupy arid regions.

The closest relative amongst the eastern leafless
species to B. walkeri is unclear. From the Riparia
subgroup, Bossiaea riparia is similar in having cladodes
with hairy faces, while B. peninsularis is similar in terms
of its striate bracts and bracteoles. However, in terms
of caducous bracts and bracteoles and numbers of
inflorescence scales, B. walkeri is closer to the Bracteosa
subgroup.

Names of uncertain application

Bossiaea humilis Meisn., in J.G.C.Lehmann,
Plantae Preissianae 1 (1): 85, adnot. (1844)

Type: [Protologue: 'Circa Sydney, ora orient., legit
Anderson, n. 78 (v. s. in Herb. Shuttleworth.)',] New South
Wales. Sydney region, Anderson 78, 1837; holotype: BM
939751, image seen MEL

The type specimen does not match any other material
seen in the course of this revision. It may be a hybrid as
it appears to be somewhat intermediate between B.
stephensonii q.v, and several other species that occur in
the Sydney region, including B. obcordota, B. nummularia
and B. prostrata.

Bossiaea linnaeoides G.Don, Gen. Hist 2:129
(1832)

Type: not designated. [Protologue: 'Native of New
Holland'.] There is insufficient information from the
protologue to identify this taxon and there is no known
type material.

Bossiaea plumosa Hort. ex Har., Diet Hort [Bois]
1(7): 195 (1893)

Type: not designated. [Protologue has no locality or
collector information.] There is insufficient information
from the protologue to identify this taxon and there is
no known type material.

Acknowledgements

I am grateful to Collections staff at the Royal Botanic
Gardens Melbourne for their assistance with mapping,
loan requests and processing, to Dr Lachlan Copeland
for making some valuable field collections, and to
AD, BRl, CANB, HO, MEL, NE and NSW for making their
collections available for study. This study was funded
by Australian Biological Resources Study (ABRS Grant
no. 207-01), which is a program within the Department
of Sustainability, Environment, Water, Population and
Communities.

References

Bentham,G. (1864).'Bo5s/oe(3'. In G.Bentham, Flora Australiensis,
Vol. 2, pp. 154-168. Lovell Reeve & Co.: London.

Lee, A.T. (1970).Taxonomic Notes on Platylobium, Bossiaea and
Templetonia in New South Wales. Contributions from the New
South Wales National Herbarium 4(3), 96-105.

Lee, A.T. (1981). Bossiaea oligosperma A. Lee, sp. nov. (Fabaceae:

Bossiaeeae). Telopea 2(2), 215-217.

Lee, A.T. and Thompson, J. (1984).'Fabaceae, Part 2' In Flora of
New South Wales. National Herbarium of New South Wales,
pp. 93-178. V.C.N.BIight, Government Printer: New South
Wales.

McDougall, K.L. (2009). Four new species related to Bossiaea
bracteosa F.Muell. ex Benth. in south-eastern Australia.
Te/opeo 12(3), 347-360.

Ross J.H. (1991). Bossiaea arenicolo (Fabaceae), a new species
from northern Queensland. Muelleria 7(3), 371 -374.

Ross, J.H. (2006). A conspectus of the Western Australian
Bossiaea species (Bossiaeeae: Fabaceae). Muelleria 23, 15-
143.

Ross, J.H. (2008). A new species of Bossiaea (Fabaceae:

Bossiaeeae) from Victoria. Muelleria 26(2), 54-56.

Thompson, I.R. (20na). A revision of Goodia (Fabaceae:

Bossiaeeae). Mue//er/a 29(2), 141-153.

Thompson, I.R. (2011b). A revision of Platylobium (Fabaceae:

Bossiaeeae). Muelleria 29(2), 154-172.

Thompson, I.R. (2011 c). A revision of Muelleranthus, Ptychosema
and Aenictophyton (Fabaceae: Bossiaeeae). Muelleria 29(2),
173-189.

Walsh, N.G. and Stajsic, V. (2007). A census of the vascular plants
of Victoria, 8"' edn. National Herbarium of Victoria, Royal
Botanic Gardens Melbourne: South Yarra.

172

Vol 30(2) 2012

Eastern Bossiaea

Appendix: Index of scientific names

Epithets of accepted names are in roman typeface, and also in
resurrected in this revision. Epithets of synonyms are in italics.

bold if the taxon is new, has new status, or has been

Name

Number ref.

Page no.

Bossiaea Vent.

alpina I.Thomps. (new species)

120

arenicola J.H.Ross

157

armitii F.Muell.

163

bombayensis K.LMcDougall

167

bossiaeoides (Benth.) A.B.Court

Introduction

107

bracteosa RMuell. ex Benth.

169

brownii Benth.

155

buxifolia A.Cunn.

133

carinalis Benth.

153

cinerea R.Br.

123

cinerea var. rigida Rod way

144

cinerea var. rosmarinifolia (Lindl.) Benth.

125

cinerea var. tenuicoulis (Graham) J.M.BIack

123

concolor (Maiden & Betche) I.Thomps. (new status)

152

coccinea Bonpl.

123

cordifolia Sweet (resurrected)

126

cordigera Benth. ex Hook.f.

130

dasycarpa I.Thomps. (new species)

140

decumbens F.Muell. (resurrected)

132

distichoclada F.Muell. (resurrected)

117

ensata Sieberex DC.

159

foliosa A.Cunn.

115

fragrans K.LMcDougall

166

grayi K.LMcDougall

168

hendersonii Regel

130

heterophyllaVent.

149

heterophyila var. stenoclada Domin

149

horizontaiis (unpublished)

130

humilis Meisn.

Uncertain appl.

172

kiamensis Benth.

127

lanceolate (Andrews) Sm.

149

lenticularis Sieberex DC.

129

ienticularis Lodd., G.Lodd. & W.Lodd.

150

linnaeoides G.Don

Uncertain appl.

172

microphylla (Sims) Sm.

145

milesiae K.LMcDougall

166

neoanglica F.Muell.

134

nummularia Endl. (resurrected)

137

obcordata (Vent.) Druce

145

obovata I.Thomps. (new species)

143

oligosperma A.T.Lee

157

ovata (Andrews) Sm.

149

Muelleria

173

Thompson

Name

Number ref.

Page no.

peninsularis l.Thomps. (new species)

162

plumosa Har.

Uncertain appl.

172

prostrata R.Br.

138

prosfram var. Tuan Creek M.S.CIemens AQ22827

140

rhombifolla Sieber ex DC

150

rhombifolia subsp. concolor (Malden & Betche) A.T.Lee

152

rhombifolia var. concolor Malden & Betche

152

riparla A.Cunn. ex Benth.

164

rosmarinifolia Lindl.

125

rotundifolia DC

150

rupicola A.Cunn. ex Benth.

154

scolopendria (Andrews) Sm.

161

scortechinil F.Muell.

142

sericea l.Thomps, (new species)

119

sp. A sensu S,W.LJacobs St J.PIckard

157

stephensonli F.Muell.

148

tasmanica l.Thomps. (resurrected, new status, new name)

144

tenuicaulis Graham

123

vombata J.H.Ross

168

walker! F.Muell.

171

Platylobium

lanceolatum Andrews

149

microphyllum Sims

145

obcordatum Vent.

145

ovatum Andrews

149

scolopendrium Andrews

161

174

Vol 30(2) 2012

Genetic analysis suggests a wide
regional provenance distribution for
Epacris impressa (Ericaceae)

Melanie Conomikes*, Gregory M Moore, Cassandra McLean^

Melbourne School of Land and Environment, University of Melbourne, Burnley Campus, SOOYarra Boulevard,
Richmond 3121.

*corresponding author: melanie.conomikes@alumni.unimelb.edu.au

Introduction

Epacris impressa Labill. is a woody heathland shrub common to parts
of south-eastern Australia. It is the floral emblem of Victoria and its
range covers most of Victoria, parts of South Australia, Tasmania and
southern New South Wales. Until recently, the strategy of'local is best'
has traditionally been accepted as best-practice for the collection of
germplasm for native plant restoration projects. However, current studies
have introduced the use of genetic fingerprinting techniques to establish
plant provenance including Random Amplified Polymorphic DNA
(RAPDs), Inter Simple Sequence Repeats (ISSR) and Amplified Fragment
Length Polymorphism (AFLP) (Bussell eta!. 2006; Krauss & Koch 2004;
Krauss etal. 2005). ISSRsand RAPDs have been used togetherto determine
phylogenetic relationships (Awasthi et al. 2004; Isshiki et al. 2008; Iruela
etai. 2002; Levt & Rowland 1997; Mattioni etal. 2002; Pharmawati et al.
2004) and assess genetic diversity (Awasthi et al. 2004; Ayres & Strong
2001; Esselman etal. 1999; Jain etal. 1999).Three collection ranges were
suggested for a 22,000 km^ area in the Sydney basin using combined
genetic and morphological assessment techniques: 1) narrow collection
range - an area as close to the planting site as possible; 2) intermediate
collection range - extending the collection area to fragmented remnants
that were once contiguous with the site; 3) regional collection range -
widening the geographic area to include a larger region (Burgin et al.
2005; Mortlock 2000). In this study, RAPDs and ISSR were performed to
determine relatedness among and within geographic sites and floral
colour races of £ impressa.

The size of heathland ecosystems has been drastically reduced and

continuesto beat riskduetocontinued land cleahng.Threatsto heathland
areas include property and pasture development, forestry, mining, fire
control in urbanised areas and infection by P/iyfop/ 7 f/)orac/nnamom/(The
State of Victoria 2002; Williams etal. 2001). Very little research has been

Abstract

Epacris impressa has showy flowers
that fall into three general colour races:
red, pink and white. It is primarily
an outcrossing species with some
examples of selfing occurring in each
population. Genetic fingerprinting
techniques were used to examine
relationships between geographic
sites and flower colour populations
and to aid provenance determination.
Results indicated that E. impressa
has a high level of genetic diversity
between and amongst sites and floral
colour races.This suggests a wide
provenance distribution for the species
which would concur with earlier
morphological studies conducted In
the1970's.

Keywords: ISSR, RAPDs, provenance,
Ericaceae, heathland

Muelleria 30(2): 175-182 (2012)

K^yal

Hotainc

Gardens

Mflbourne

Muelleria

175

Conomikes et al.

Figure 1. £ impressa falls into three general floral colour races: a) red (Angahook-Lorne Site A); b) pink (Cranbourne Site CA) and c)

white (Cranbourne Site C).

conducted into the genetic provenance of plant species
in these shrinking areas of heathland and provenance
is often assigned by local anecdotal information rather
than on the basis of genetic traits.

A major study by Stace and Fripp in 1977 described
polymorphism in £ impressa (Stace & Fripp 1977a,
1977b, 1977c). The flowers of £ impressa fall into three
general races: red (fuchsia/deep pink), pink (light pink)
and white (Fig. 1). Populations of £ impressa are often
polymorphic for flower colour with monomorphic
populations occurring less frequently (Stace & Fripp
1977c). Polymorphic populations exist geographically
between monomorphic populations and appear to
provide a conduit for gene flow (Stace & Fripp 1977b).
Soil pH has been shown not to be a determining factor
in flower colour and it is possible that gene flow is
influenced by vector selection (Stace & Fripp 1977b,
1977c). Flowering times of different flower colours
overlap at polymorphic sites but are staggered between
locations by flower colour in monomorphic populations
(Stace & Fripp 1977b).

Polymorphism could be an ancient trait present in
£ impressa or a more recent development caused by
hybridization between populations (Stace & Fripp 1977a,
1977c).Theconsistencies between corolla colour, length,
and anther colour led Stace and Fripp (1977a, 1977b) to
postulate that raciation in the species may be an early
indicator of species division. However, further work by
Fripp (1982) concluded that £ impressa is primarily an
outcrossing species with no disparity between seedset
or viability when races were crossed. Fripp (1982) self-
pollinated and cross-pollinated individuals of different

races from a variety of populations over a five-year
period. Morphological study of germinated plants
indicated that the diverse races were not separate
species (Fripp 1982). The evidence suggested that
raciation was probably a long-standing trait rather than
an early sign of species divergence. This correlates with
later genetic research that included the genus Epacris in
Ericaceae due to early genetic links. This research was
the first to use the Inter Simple Sequence Repeat (ISSR)
method with a member of the Southern Hemisphere
Ericaceae.

Methods

Six research sites with remnant populations of £ impressa
were selected based on their geographic separation
and to provide examples of both polymorphic and
monomorphic flower-colour demes. The research sites
used for genetic studies were Angahook-Lorne State
Park, Victoria (Site A) (38°37'S, 143°53'E), Braeside Park
Heathland (Site B), Victoria (37“59'S, 145’08T), two sites
at the Royal Botanic Gardens Cranbourne, Victoria (sites
C and CA) (38"07'S, 145“16'E and 38“08'S, 145‘15'E),
the Grampians National Park, Victoria (site G), Sundial
track (37'’10'S, 142®30'E), and a privately-owned site in
Tullah, Tasmania (siteT) (41*44'$, 145*37'E). No white-
flowered plants were observed at Angahook-Lorne
State Park during the collection period and Braeside
Park Heathland was monomorphic for white-flowered
plants.

Genetic fingerprinting using RAPDs and ISSR was
conducted in two experiments with samples from each

176

Vol 30(2) 2012

Epacris impressa colour races and provenance

of the three flower colour races (white, pink and red)
found at each site. After initial primer screening, ISSRs
and RAPDs were performed on a subset of red, pink
and white samples from each site using primers that
produced the clearest reproducible banding patterns as
per the following protocols.

Cuttings consisting of 10-20 cm of new tip growth
and flowers were collected from flowering E impresso
plants from Angahook-Lorne State Park and the
RBG Cranbourne sites in September 2001, from the
Grampians in November 2001 and from Braeside Park
Heathland and Tullah, Tasmania in September 2002.
Each sample was given an alphanumeric identification
according to site (A = Angahook-Lorne State Park, B =
Braeside Heathland, C and CA = Royal Botanic Gardens
Cranbourne sites, G = Grampians National Park, T =
Tullah, Tasmania), colour (W = White, R = Red, or P =
Pink) and the plant of origin. Flower colours were further
identified using the Royal Horticultural Society Colour
Chart (cl 995). Plant material collected at each site was
stored at -20‘‘C until required (Table 1). Herbarium
specimens were lodged at the National Herbarium of
Victoria (MEL) after DNA isolation from material.

DNA was isolated from the £ impressa flowers
using the Qiagen DNeasy® Plant kit according to the
manufacturer's instructions. Electrophoresis was
performed on a Horizon'58 Life Technologies™ Gibco
BRL Horizontal Gel Electrophoresis Apparatus gel tray
with 1% agarose inTBE stained with ethidium bromide
and run at 96 V for 30 minutes. Bands of DNA were
visualised on a UV transilluminator and photographed

Table 1 . Accession numbers for herbarium vouchers. An
indicative sample was submitted for each flower colour
race for sites G (Grampians National Park), CA (Royal Botanic
Gardens Cranbourne), and A (Angahook-Lorne State Park).
All plant material was used for DNA isolation from samples
collected at sites B (Braeside Heathland), C (Royal Botanic
Gardens Cranbourne), and T (Tullah, Tasmania). No white-

flowered plants were observed at Angahook-Lorne State Park
during the collection period.

MEL Accession numbers

Site

Red/Fuchsia

Pink

White

2337596

2337597

2337594

2337595

2337593

2337591

2337592

2337590

with a Kodak Digital Science DC120 digital camera. The
RAPD OPB primer set was found to be successful with
the ericoid, Vaccinium macrocarpon Ait. (Stewart & Nilsen
1995) and was selected for screening with £ impressa.
The Operon OPB set of 20 primers (OPB1 GTTTCGCTCC,
OPB2 TGATCCCTGG, OPB3 CATCCCCCTG, OPB4
GGACTGGAGT, OPB5TGCGCCCTTC, OPB6TGCTCTGCCC,
OPB7 GGTGACGCAG, OPB8 GTCCACACGG, OPB9

TGGGGGACTC,

OPB10

CTGCTGGGAC,

OPBII

GTAGACCCGT,

OPB12

CCTTGACGCA,

OPB13

TTCCCCCGCT,

OPB14

TCCGCTCTGG,

OPB15

GGAGGGTGTT,

OPB16

TTTGCCCGGA,

OPB17

AGGGAACGAG,

OPB18

CCACAGCAGT

OPB19

ACCCCCGAAG) was screened initially with two DNA
samples, one a red-flowered plant from Angahook-Lorne
State Park (site A) and the other a white-flowered plant
from Cranbourne (site C), using the protocol outlined
below. Each sample and primer was run twice to test for
reproducibility. Bands were obtained for both samples
with the RAPD primers OPB1, OPB3, OPB4, OPB5, OPB6,
OPB7, OPB15, and OPB19. From these, the two primers
that yielded the clearest reproducible bands, OPB6 and
OPB19, were chosen for RAPD experiments

Each RAPD/PCR reaction consisted of the following:
3.2 pi 1.25 mM dNTP, 0.8 pi 25 mM MgCI^ 2 pi Tris-HCl
reaction buffer, 12 pi milli-q Hp, 1 pi Operon OPB
series sequence primer, and 0.5 p! QIAGEN Tag DNA
Polymerase. The RAPD reactions were run in a Biometra*
Personal Cycler top-heating thermocycler using a 4-
minute strand separation cycle at 94°C, then 45 cycles
of one minute at 94°C, two minutes at 36^, and two
minutes at 72°C and a final 72“C extension step for 10
minutes. A negative control with no DNA was included
in each run to test for contamination. Gels were run as
previously described.

Six standard ISSR primers, 812 (GAG AGA GAG AGA
GAG AA), 814 (CTC TCT CTC TCT CTC TA), 824 (TCT CTC
TCTCTCTCTCG), 835 (AGA GAG AGA GAG AGA GYC), 836
(AGA GAG AGA GAG AGA GYA) and 857 (ACA CAC ACA
CAC ACA CYG) were screened with two DNA samples as
for RAPD screening.The primers were selected based on
their ability to amplify loci in a variety of plant genera
and species (Casasoli et al. 2001; Ge ef al. 2003; Levi &
Rowland 1997; Mattioni ef al. 2002; Nan ef ol. 2003;
Pharmawati et al. 2004; Qiu ef al. 2004; Wang ef ol. 2004;
Xiao & Berch 1996). ISSR reactions were prepared as

Muelleria

177

Conomikes etal.

Table 2. Similarity matrix for the entire sample set using the Jaccard coefficient. A = Angahook-Lorne State Park, B = Braeside
Heathland, C and CA = Royal Botanic Gardens Cranbourne sites, G = Grampians National Park,T=Tullah,Tasmania.The site letters
are followed by a letter indicating floral colour race W (white), R (red) or P (pink) and sample number.

BWI

031

OV2

CR CR
133 134

lot

105

121

W24

W27

W34

CAR

CAP

118

(;\\

119

I2U

137

110

127 TWl

ru3

TRl

TR2

TPI TP2

TP4

ARI8

1.00

ARI9

0.75

1.00

AP34

0.00

1.00

Bwn

0.00

1.00

BWI2

0.00

0.57

1.00

C\V19

0.33

0.38

0.29

0.18

oox

1 00

O\20

0.^5

0.29

0.40

0.22

020

058

100

CRI33

025

0.13

0.00

0.33

031

0.27

0.21

1.00

CRI34

0.29

0.14

0.25

0.15

0.14

0.29

0.23

0.75 LOO

CPIOl

0.18

0.20

0.13

0.29

0.42

0,36

0.20 0.10

1.00

CPI05

0.13

0.14

0.33

0.17

040

0.30

0.38

0.11 O.OU

0.56

1.00

CP12I

0.09

0.10

0.25

0.13

0.29

0.33

0.40

0.20 0.10

0.89

0,63

1.00

CAW24

0.00

000

033

000

DOQ

0.27

020

0.21 0 33

0.14

0.00

0.14

1.00

CAW27

0.00

000

0.33

000

0.09

0.27

020

0.21 033

0.14

0.00

0.14

0.71

LOO

CAW34

0.00

flJ3

0.00

O.IQ

0.18

0.10

0.23 036

0.00

0.57

0.83

1.00

CAR4

0.20

0.25

0.20

0.14

0.29

O.IS

0.09

0.08 0.18

U.IO

0,00

0.00

0 13

0.29

0.33

LOO

TAPIS

0.20

050

0J5

a2S

0 29

0.33

0.67 0 31

0.50

060

0.57

000

0.00

000

0.00

1.00

CAPI7

0.25

0.50

033

0.33

040

0.33 0 00

0 38

0.75

0.43

000

0.75

LOO

C\VI18

000

0.00

0 25

0 20

0.17

0.20

0.25

0 29 0 14

0.22

000

0,25

0,14

0.14

0.00

0,17

0,00

1.00

CW 119

000

0.00

025

0 20

D.I7

0.20

0,25

aJ9 0 14

0.22

0.00

0.25

0.14

0.00

ouo

0.17

0.00

1.00

LOO

(:W120

0.00

025

0.20

0.17

0.20

0.25

0.29 0 14

0.22

U.OO

0 25

0.14

O.IW

0.00

0,17

000

1.00

1 00

LOO

(:RI37

0.13

0.20

0.25

0.22

0.29

0 36

0.45 0.36

0.25

0.10

027

0 1(1

O.IO

0.11

000

040

0 20

033

0,33

LOO

CPIlO

0.00

0.33

O.DU

0.00

0.27

0.20

0.25 0.50

0 38

0,17

0 43

029

0 29

a3J

0 tl

040

0 20

O.lt

0.43

1 00

CPI27

ODO

0.00

0 33

000

ono

0 08

0.09

025 0 50

0.22

000

0.25

0 29

0.29

033

0.11

017

UOO

0.25

0 50

LOO

TWl

0.25

020

0.1?

0.33

043

0 29 014

0.38

0.40

0.43

000

(1,75

0.50

0J3

0.33

0,60

0.25

0.11 LOO

T33'3

0.25

0.20

0.25

0.20

0.33

0.43

0,14 0,00

0.38

040

0.43

UCW

000

0.00

0.40

0.50

0.14

0.60

0.25

0.11 0.60

LOO

IRl

0.00

0J3

0.17

0.30

0.22

0,43 0.29

0 40

0.43

0.44

0 20

020

a25

0.00

060

0 40

0 17

0.17

0.38

0.40

0.17 0.40

0.17

1.00

IR2

0.00

u.oo

I.WI

0.17

0,18

0.10

0.43 0.29

0.27

0.25

0,30

0.20

0 20

0.25

0,00

060

0 40

0.40

D.40

0.38

0,40

0,40 0,40

0.17

0,67

LOO

TPI

0.50

0|7

000

0.14

05<S

033

O.ll 0.13

0.50

0J3

0,38

ni3

0.29

0.»4

0.25

0.33

0.4(1

o.n

0.11

O-ll

0.11

020

0.00 0.2S

0.25

0.14

LOO

TP2

0.50

0.25

0.00

0.10

0.55

0.36

0.18 0.33

0.50

0.33

0.38

0.33

0.50

0.38

0.30

0.33

0.40

0.08

008

0.08

0.36

0.25 0 18

0.18

0.14

0.67 1.00

TP4

0.50

0.20

0.00

0.63

0.38

0.00 0.00

0.50

0.33

0.38

0.14

0.00

0.13

0.33

0.40

0.13

0.22

0,00 0.29

0.29

0.14

0.83 0.56

LOO

for RAPDs and run with an initial three-minute cycle at
94°C, then 45 cycles of one minute at 94°C, 45 seconds
at 54°C, one minute at 72®C then a final extension step
of 10 minutes at 72°C

Bands were obtained for both samples with ISSR
primers 812, 824, 835, and 836. From these, the two
primers that yielded the clearest reproducible bands,
812 and 836, were chosen for ISSR experiments as per
previously described protocols. Gel photos were visually
reviewed and faint ambiguous bands were removed
from the data set (Williams et al. 1990; Zawko et al,
2001). All of the gel photos were reviewed with their
corresponding loci information from the KodakID
analysis program. High (1500,2000 and 3000 base pairs)
and low (100, 200, 300 base pairs) molecular weight
bands were also removed from the analysis to minimise
the possibility of the inclusion of nested inverted repeats
(Stewart & Excoffier 1996; Stewart & Nilsen 1995). Any
individuals with no bands were excluded from the data
analysis. Monomorphic bands were not present in any
of the RAPD or ISSR results. A total of 182 polymorphic
bands were scored with four primers for 31 individuals.

The molecular weights were then converted into
binary data for presence (1) or absence (0) of bands.

Any samples that were missing data from more than
one primer were removed and the Exeter software
program NTSYSpc 2.20e was used for statistical
analyses. A similarity matrix was generated from
the data set using the Jaccard coefficient, a/(n-d)
(Table 2). Reduced similarity matrices were created with
data sub-sets by site (Table 3) and floral colour race
(Table 4) to compare the average similarities between
and amongst populations. A dendrogram was created
using a clustering algorithm with an unweighted pair-
group method, arithmetic average (UPGMA) formula to
illustrate relationships between individuals in the entire
sample group (Fig. 2).

Results

ISSR and RAPD analyses showed a high level of genetic
variability both between and amongst geographic
and race populations of £ impressa. Individuals did not
cluster (Fig. 2) by site or flower colour (except for small
clusters of 2-3 individuals from the same site and flower
colour) and inter-site similarity coefficients ranged from
13-30% and inter-colour similarity coefficients from
19-31%, with intra-site coefficients similar. The results

178

Vol 30(2) 2012

Epacris impressa colour races and provenance

-AR18
-AR19
-TP1
-TP4
-TP2
-CW19
-CW20
jAP34
TR2
-TR1
-CPI 01
-CP121
-CP105
-CAP15
-CAP17
-GR137

-TW1

-TW3

-CAW24

-CAW27

-CAW34

-CR133

-CR134

-GP110

-GP127

GW118

-GW119

GW120

-BW11

-BW12

-CAR4

0.36

Cocfllcicnl

Figure 2. Dendrogram illustrating phylogenetic relationships between individuals in the entire population sample. The first
letter or letters represent the collection site (A, B, C, CA, G, T) and the second letter the flower colour race {R, P, W). The number

corresponds to the plant identification number.

Muelleria

179

Conomikes etoL

Table 3. Reduced similarity matrix showing average
percentages of similarity by site. A = Angahook-Lorne State
Park, B = Braeside Heathland, C and CA = Royal Botanic
Gardens Cranbourne sites, G = Grampians National Park,T =
Tullah, Tasmania.

SITE

0.13

0.00*

0.19

0.22

0.21

0.26

0.20

0.15

0.24

0.17

0.10

0.13

0.23

0.14

0.30

0.12

0.33

0.25

0.23

0.24

* white-flowered plants were not found at site A at the time of
collection for DNA isolation.

indicated a 20% average genetic similarity between
geographic populations and a 22% average similarity
between colour races. The pink-flowered race had the
highest average intra-population polymorphic similarity
at 31 %,The red and white races had the lowest inter-race
and intra-race average genetic similarities (19-23%).

The percentages of similarity ranged from 10%
between sites A and G to a 33% average possibility of
relatedness between individuals at sites C and T (Table
3). Site A had the lowest percentage of genetic similarity
(13%) of plants within its population, followed by sites
CA (17%) and B (19%). Site G had the highest amount
of intra-site genetic similarity (30%) followed by site
C (26%) and site T (24%). There was a 20% inter-site
average polymorphic similarity between plants.

Plants of the pink-flowered race had the highest
average percentage (31 %) of intra-race genetic similarity
(Table 4). White-flowered and red-flowered plants had
lower average probabilities of being genetically similar
to other plants of the same flower-colour race (21%
and 19% respectively). Pink-flowered plants had a 21%
average genetic similarity to white-flowered plants and
a 23% similarity to the red-flowered race. Red-flowered
plants had an equal average probability (19%) of being
related to either other red-flowered or to white-flowered
plants. There was a 22% average genetic similarity
between plants of any one flower-colour to those of any
other race.

Clustering can be seen between individuals of the
same flower colour race within populations, with
the Grampians white-flowered plants (GWl 18-120)

Table 4. Reduced similarity matrix showing average
percentages of similarity by floral colour race
(W = white, R = red, P = pink).

0.21

0.19

0.21

0.23

0.31

showing the highest similarity coefficient (Fig. 2).
Individuals from each site are primarily clustered at
low similarity coefficients or are seen grouped with
members of other site populations indicating a high
level of genetic variability. Clustering at the highest
similarity coefficients is shown primarily by small but
separate groups of red, pink, and white flowered plants.
The smaller discreet colour alignments, rather than
three major groupings by floral race, suggest a relatively
high level of genetic variability within floral races and
within sites.

Discussion

During this research and previous studies, (Stace & Fripp
1977a) plants of different flower colours in populations
that were polymorphic for flower colour were observed
to have separate but overlapping periods of flowering,
which would have an influence on the genetic
composition of the separate floral races. Native and
introduced vectors that select by flower colour during
overlapping periods of flowering of the three floral
races would not be cross-pollinating between the floral
races (Castellanos ef al. 2003; Melendez-Ackerman et
al. 1997; Streisfeld & Kohn 2006). Differences in the
time of flowering between flower-colours in the same
population could also have an influence on pollination
patterns. Research is currently being undertaken
at LaTrobe University on pollinator flower-colour
preference with £ impressa (Webster N., Edwards
T. & Hoebee S. pers. comm. 2012). This may provide
further insight into vector influence on flower colour
composition of polymorphic populations.

The geographic isolation of the Grampians has
created many unique taxa and it was not surprising
that the Grampians population (site G) showed the
highest level of intra-site genetic similarity (30%). A 26%
average polymorphic similarity between individuals

180

Vol 30(2) 2012

Epochs impressa colour races and provenance

was recorded at the Royal Botanic Gardens, Cranbourne,
site C. This site primarily contained plants of the white-
flowered race, which may account for the high level of
intra-site genetic similarity. The Tullah site in Tasmania
demonstrated a 24% average intra-site similarity. The
site consisted of an area of remnant vegetation not
directly geographically linked to other populations.

Angahook-Lorne site A had primarily red and pink
flowered populations, and demonstrated the lowest
average genetic similarity within its own population
(13%). This concurs with Stace and Fripp's (1977b)
findings of high levels of polymorphism for flower-
colour race within linked 'mosaics' of populations. The
site was also part of a large coastal range of linked
E, impressa populations within a protected state park
region. This has provided the opportunity for genetic
dispersal to other geographically linked populations
not sampled. Genetically, the site appears to be part of a
larger population or one of a series of linked populations
that share genetic traits. The recent creation of the
Otways National Park will help to preserve the genetic
integrity of this large polymorphic group.

Braeside site B and Cranbourne site CA also had low
average intra-site polymorphic similarities at 13% and
17% respectively. These results were surprising since
both sites were geographically removed from other
populations. Site B comprised all white-flower race
plants which could explain the polymorphic similarities.
Site CA had a robust mix of races but all E impressa
plants at the site died during this research due to long¬
term drought conditions.

Epacris impressa appears to have a large regional
provenance distribution. These findings concur with
Stace and Fripp's (1977a; 1977b; 1977c) earlier work
on raciation of the species which found high levels of
polymorphism for flower colour race, corolla colour and
corolla length. While geographic populations showed
high levels of inter-site genetic diversity, morphological
characteristics should still be considered when
collecting propagative material for revegetation. Since
genetic fingerprinting techniques target unknown
regions of the genome, locally adaptive traits may not be
represented by the loci in RAPD and ISSR analysis (O'Brien
etal. 2007). A combination of genetic fingerprinting and
traditional morphologic observation is recommended
to determine provenance (Krauss etai. 2005) and plant
propagules should still be collected from populations of

similar floral colour race (Stace & Fripp 1977b). Hence,
when trying to re-establish a population of all white-
flowered plants, it is suggested that cutting material is
collected from another site of all white-flowered plants.
Geographic proximity of populations does not appear
to be an issue in provenance determination based
on the regional genetic spread of E impressa. This has
been found to be the case in other Australian studies
where large regional areas of provenance have been
established with DNA fingerprinting techniques (Krauss
& Koch 2004; Krauss etal. 2005).

Based on this research, £ impressa has a high level
of both intra-race and inter-race genetic diversity. The
inter-site genetic similarities indicate that E impressa
populations have probably been geographically inter¬
connected until recently. Further work will be required
on Tasmanian populations to determine their genetic
links to mainland South-eastern Australia plants and
additional studies in southern New South Wales and
South Australia would be valuable.

Acknowledgements

Our thanks to the curatorial staff at the National
Herbarium of Victoria, editors and reviewers for
comments on this paper. Sincere thanks and
appreciation to a dedicated researcher and mentor.
Dr Cassandra McLean', who succumbed after a long
and hard battle with cancer in 2009. Thank you to the
Australian Flora Foundation for a generous grant that
helped support this research.

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182

Vol 30(2) 2012

Cyclosorus interruptus
(Thelypteridaceae): new to Victoria

Steve Sinclair\ Val Stajsic*^and Geoff Sutter^

’ Arthur Rylah Institute for Environmental Research, Department of Sustainability and Environment. 123 Brown St,
Heidelberg, Victoria, 3084.

^ National Herbarium of Victoria, Royai Botanic Gardens Melbourne. Private Bag 2000 Birdwood Avenue,

South Yarra 3141.

^Corresponding author val.stajsic(5)rbg.vic.gov.au

Introduction

Abstract

During recent botanical survey work in south-western Victoria {Sinclair
and Sutter 2008), a fern not referrable to any species recorded previously
in Victoria was encountered (Fig. 1). Examination of the material
confirmed the identity ofthespecies as Cyc/osorus/>7ferrupfus(Willd.)H.lto
(Thelypteridaceae). This species occurs in the tropics and sub-tropics of
all continents as well as New Zealand (Bostock 1998). In Australia, prior to
the current discovery, it was known to occur in tropical central Australia,
and southward along the margins of the continent about as far south as
Sydney and Perth.

In Victoria the family Thelypteridaceae is represented by four species
in four genera: Christella dentata (Forssk.) Brownsey & Jermy, Cyclosorus
interruptus (Willd.) H.lto (here reported), Pneumatopteris pennigera
(G.Forst.) Hoittum, and Thelypteris confJuens (Thunb.) C.V.Morton.
Cyclosorus interruptus can be distinguished from Christella dentata
and Pneumatopteris pennigera by the presence of scattered, papery,
broad, flat scales on the pinnae midribs on the lower surface of mature
fronds (Fig. 2a), which are absent in Christella and Pneumatopteris;
similar scales are present in Thelypteris confluens, and pale brown ovate
scales are sometimes present on the lower surface of young fronds in
Pneumatoperis pennigera. The upper surfaces of the pinnae in Cyclosorus
interruptus are virtually hairless (occasional, minute, pointed hairs
on veins present), whereas in Christella dentata the upper surface of
pinnae have many short, pointed hairs. Cyclosorus interruptus also has
stalkless, spherical orange or orange-red glands on the veins on the
lower surface of the fronds, which are absent in Christella dentata and
Pneumatopteris pennigera (Fig. 2b).The texture of the fronds of Cyc/osorus
interruptus is harsh, whereas the fronds of Pneumatopteris pennigera
are softer-textured. When sori are present, Cyclosorus interruptus is
easily distinguished from Pneumatopteris pennigera by its indusiate (i.e.
protected) sori; the sori of P. pennigera lack indusia.The absence of sori

A new fern, Cyclosorus interruptus, is
reported for Victoria. Information is
provided as to how this species can
be distinguished from closely-related
Victorian ferns. Its Victorian distribution
is discussed, along with Its habitat,
the threats to its persistence and its
conservation significance in Victoria.

Key words: Ferns, Cyclosorus,
identification, ecology, Australia.

Muelleria 30(2): 183-188 (2012)

!<uyal

Uotatlic

CJardens

Mclhourt>«

Muelleria

183

Sinclair efcj/.

on the lowermost 1 (-2) of the basal pair of veins (the
pair that unite below the sinus) is also diagnostic. The
sori are usually present in the corresponding position
in Christella dentata and Pneumatopteris pennigera.
Cydosorus interruptus differs from Thelypteris in that the
pinnae are lobed for about one-half to one-third of the
distance to the pinnae midribs, with the basal veins in
adjacent pinnae lobes always branching to produce
a long excurrent vein passing to the sinus membrane,

Figure 1. Cydosorus interruptus in situ.

Figure 2. Frond under-surfaces, showing diagnostic features.
A: Cydosorus interruptus, with the broad scales indicated
(absent in Pneumatopteris and Christella), and the pinnae
division clearly evident; B: C interruptus with the sessile glands
highlighted; C: Thelypteris confluens, showing the absence of
glands, the presence of scales and the relatively deeper pinnae
division.

whereas in Thelypteris the pinnae are lobed almost to
the pinnae midribs, and all veins are free (Bostock 1998)
(Fig. 2c).

Many ferns are readily dispersible and popular in
cultivation, making it potentially difficult to determine
whether some species are historically indigenous
to a given area (e.g Pteris umbrosa at sites such as
Yellingbo, Victoria, distant from its conventionally
recognised distribution in eastern Victoria, Stajsic

184

Vol 30(2) 2012

Cydosorus interruptus in Victoria

pers. obs.). We consider Cydosorus interruptus to be
indigenous in Victoria, since the species is apparently
very rarely (although easily) cultivated (on the basis of
internet searches and published literature, e.g. Jones
& Clemesha 1993), the individual plants appear long-
established, and the area of occurrence is sparsely
inhabited (although two farmhouses are within 1 km).
The site is unfenced and the possibility that it may have
been introduced with stock, although remote, cannot

be discounted. However, given the remarkable disjunct
Australian occurrence of Thelypteris confJuens in north¬
eastern Victoria, otherwise known with certainty only in
south-eastern Queensland, the occurrence of Cyc/osorus
interruptus in south-western Victoria is perhaps less
surprising given the far greater natural range of this
species in Australia. As is often the case with Cydosorus
interruptus, Thelypteris confJuens has a preference for
swampy habitats (Wilson 1990; Bostock 1998).

Figure 3.The Victorian
habitat of Cydosorus
interruptus when A. dry; and
B. inundated.

Muelleria

185

Sinclair era/.

Table 1. Species associated with Cyclosorus interruptus in Victoria, taken from two quadrats (D0076200, D0076300).The
abundance values are consistent with the Victorian Flora Site Database (2007), where the information from these quadrats is
stored.The species are listed by their abundance, then alphabetically. The nomenclature for botanical names follows Walsh &
Stajsic (2007). Vernacular names follow the Victorian Flora Site Database.

Species

Common name

D0076200

D0076300

Leptospermum lanigerum

WoollyTea-tree

Cyclosorus interruptus

Swamp Shield-fern

Ranunculus sp.

Buttercup

Rumex bidens

Mud Dock

Stellaria angustifolia

Swamp Starwort

Urtica incisa

Scrub Nettle

^Sonchus asper

Rough Sow-thistle

Carex appressa

Tall Sedge

Crassula helmsii

Swamp Crassula

Eleocharis acuta

Common Spike-sedge

Hydrocotyle sibthorpioides

Shining Pennywort

Persicaria decipiens

Slender Knotweed

Triglochin alcockiae

Southern Water-ribbons

Hydrocotyle muscosa

Mossy Pennywort

*Paspolum distichum

Water Couch

Glyceria australis

Australian Sweet-grass

*Nasturtium officinale

Watercress

Poa labillardierei subsp. labillardierei

Common Tussock-grass

Juncus procerus

Tall Rush

Lachnagrostis filiformis

Common Blown-grass

Leptinella reptans

Creeping Cotula

*Aster subulatus

Aster-weed

Baumea articulata

Jointed Twig-sedge

Calystegia sepium subsp. roseata

Large Bindweed

?Nasturtium microphyllum

Brown Watercress

Carex fascicularis

Tassel Sedge

Xuscuta suaveolens

Fringed Dodder

Xynodon dactylon vor. dactylon

Couch

Lilaeopsis polyantha

Australian Lilaeopsis

*Rumex conglomeratus

Clustered Dock

Xirsium vulgare

Spear Thistle

*Rumex crispus

Curled Dock

Triglochin procera

Water Ribbons

-1-

Alternanthera denticulate

Lesser Joy weed

Asperula conferta

Common Woodruff

-1-

Carex gaudichaudiana

Fen Sedge

-1-

Dichondra repens

Kidney-weed

Lobelia pedunculate

Matted Pratia

*Solanum nigrum

Black Nightshade

Solanum sp.

Kangaroo Apple

186

Vol 30(2) 2012

Cyclosorus interruptus in Victoria

Habitat and threats

The Victorian plants grow along the flats of Darlot
Creek, near Tyrendarra. Interestingly, the Tyrendarra
population of Cyclosorus occurs within a distance
of ca. n km from the similarly rare and restricted
Pneumatopteris pennigero. The surrounding landscape
at the site consists of weathered calcareous dunes,
however Cyclosorus grows on alluvial deposits of silt/
clay. The habitat is open, with occasional patches of
Leptospermum lanigerum (nearby but not directly
associated), and on some occasions is subject to partial
shallow inundation (Fig. 3). Two floristic quadrats
were taken around patches of Cyclosorus, in order to
characterise its habitat (Table 1).

Cyclosorus is long-rhizomatous, and it is difficult to
determine the number of individual plants that make up
the Victorian population without genetic analysis. We
counted 42 fairly distinct clumps, some of which cover
several square metres. These are distributed along ca.
400 m of creek-line.

Livestock presumably pose a long-term threat
to this species. The streamside habitat is unfenced
and accessible to stock (currently sheep). It appears,
however, that this species has tolerated stock for many
years, and is probably secure in the immediate-short
term in Victoria if the current management doesn't
change.

Given that the plants grow about 2 km from the coast
(less than 5 m above sea level) with obviously estuarine
elements nearby (e.g Juncuskraussii occurs in extensive
beds shortly downstream), the Victorian population
of Cyclosorus is potentially at risk from rising sea levels
which may occur as a result of climate change. We do
not have direct evidence for the tolerance of Cyclosorus
of saline conditions; however it would seem that this
species is tolerant of brackish conditions. Vegetation
studiesfrom other states show that Cyc/osorus/nferrupfus
frequently occurs in brackish-saline, estuarine or
near-coastal areas, often in paperbark swamps (e.g.,
Melaleuca quinquenervia, Kingston etal. 2004). In New
Zealand it grows near thermal springs (Bostock 1998).
Unpublished data from salinity tests over several
seasons show that Darlot Creek, including waters in
the vicinity of the Cyclosorus plants, is generally slightly
brackish (often EC ca 2 dS/m) and of neutral pH (in the

range 6.5-7.5) (J. Macdonald, Arthur Rylah Institute,
pers. comm.). Measurements on two soil samples (ca
5-15 cm depth) and a surface water sample taken from
among the Cyclosorus plants in May 2008 support this
(pH 6.S-7.7; EC 0.6-1.2 dS/m). It remains to be seen how
much salinity Cyclosorus can tolerate.

Weed invasion may also present a threat to Cyclosorus,
but probably not in the immediate future. Currently, the
abundance of weeds is relatively low in the area where
Cyclosorus occurs. Furthermore, the long-lived, strongly
rhizomatous habit of the plant might make established
plants resilient to the effects of some competition.
Weeds may, however, in future alter the site to the
extent that the germination of new plants is suppressed.

It would seem that the most pressing threat to the
persistence of Cyclosorus is its very small population size
and area of occupation, making it highly vulnerable to
extinction from chance events. Presumably, the small
and isolated Victorian population has low genetic
diversity, reducing its ability to adapt, and increasing its
vulnerability to environmental change.

Assuming that the species is indigenous to Victoria
(which we assume to be the case), we recommend that
it be assigned a Victorian conservation status of Critically
Endangered, using lUCN criteria. In the standard
notation of the lUCN Red List (lUCN, 2001): CR B1 ab(i,ii,iii
,v)+2ab(i,ii,iii,v);Cl-f-2a(i,ii); D.The National Herbarium of
Victoria recently classified the species as 'endangered' in
Victoria (Walsh & Stajsic 2007).

Acknowledgements

Am Tolsma (Department of Sustainability and
Environment) provided field assistance. David Cameron
(DSE) assisted with the lUCN assessment. We are grateful
to Dallas Mitchell (Tyrendarra, Victoria) for allowing
access to the site, and to Peter Bostock (Queensland
Herbarium) for confirming the determination of the
material. We also thank the two anonymous reviewers
for helpful comments.

References

Bostock, P.D. (1998). Thelypteridaceae. In Flora of Australia
48, 49-53. Australian Biological Resources Study/CSIRO
Publishing.

lUCN (2001). lUCN Red List categories and criteria, Version 3.1.
lUCN Species Survival Commission. lUCN, Gland, Switzerland
and Cambridge, UK.

Muelleria

187

Sinclair eta/.

Jones, D.C. and Clemesha, S.C. (1993). Australian Ferns and Fern
Allies. The Currawong Press: Chatswood.

Kingston, M.B., Turnbull, J.W. and Hall, P.W. (2004). Tweed
vegetation management strategy, 2004. Ecograph Ecological
and Geographic Information Systems Consultants,
Murwillumbah, The Tweed Council. Appendices (vol. 3).
www.tweed.nsw.gov.au. (accessed 16 July 2008).

Sinclair, S J. and Sutter, G.R. (2008). Estuarine wetland vegetation
mapping, Gleneig Hopkins CMA. Arthur Rylah Institute for
Environmental Research Technical Report Series no 178.
Department of Sustainability and Environment: Heidelberg,
Victoria.

Victorian Flora Site Database (2007). The State of Victoria,
Department of Sustainability and Environment (accessed 2
February 2012 via the 'Flora Information System', (updated
2007] -©Viridans Biological Databases).

Walsh, N.G. and Siajsic, V. (2007). A census of the vascular plants
of Victoria (Addendum). Royal Botanic Gardens Melbourne:
South Yarra

Wilson, P. (1990). Thelypteridaceae. In Harden, G.J. (Ed) Flora
of New South Wales 1. New South Wales University Press:
Kensington.

188

Vol 30(2) 2012

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Allan, H.H. (1961). Flora of New Zealand. R.E. Owen, Government Printer: Wellington.

Ahti,T. and Kashiwadani, H. (1984).'The lichen genera Cladia, Cladina and Cladonia in southern Chile'. In H. Inoue
(ed.), Studies on the cryptogams of southern Chile, pp. 125-149. Kenseisha Ltd: Tokyo.

Brummitt, R.K. and Powell, C.E. (1992). Authors of plant names. Royal Botanic Gardens: Kew.

Jarman, SJ. (1975). Experimental foxonomy on the family Epacridaceae. PhD thesis. The University of Tasmania, Hobart:
Tasmania.

Pickett, J.W., Smith, N., Bishop, P-M.. Hill, R.S., Macphail, M.K. and Holmes, W.B.K. (1990). A stratigraphic evaluation of
Ettingshausen's New England Tertiary plant locaWties. Australian Journal of Earth Sciences 37,293-303.

Swofford, D.L. (2000). PAUP. Phylogenetic analysis using parsimony (version 3.1.1.) Illinois Natural History Survey:
Champaign, USA.

Walsh, N.G. (1994).'Poaceae'. In N.G. Walsh andT.J. Entwisle (eds). Flora of Victoria, Vol.2, pp. 356-627. Inkata Press:
Melbourne.

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Royal Botanic Gardens Melbourne
Birdwood Avenue, South Yarra
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www.rbg.vic.gov.au ISSN 0077-1813

Contents

Volume 30(2) 2012

Contributed papers Page

Five new endemic eucalypts for Victoria.83

- K. Rule

A revision of eastern Australian Bossiaea {Fabaceae: Bossiaeeae).106

- IR. Thompson

Genetic analysis suggests a wide regional provenance distribution for Epacrisimpressa .175

- M. Conomikes, GM. Moore, C. McLean

Cycfosorus interruptus (Thelypteridaceae): riew to Victoria.183

- S. Sinclair, V. Stajsic and G. Sutter

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